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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

us ISShi 0027.4100

But Lett n OF THE

Museum of

Comparative

Zoology

Pleistocene AAustelidae (Mammalia,
Carnivora) from Fairbanks, Alaska

ELAINE ANDERSON

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 1
14 APRIL 1977

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dae (Mollusca: Bivalvia). 265 pp.

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chusetts, 02138, U.S.A.

PLEISTOCENE MUSTELIDAE (MAMMALIA, CARNIVORA)
FROM FAIRBANKS, ALASKA

ELAINE ANDERSON!

Abstract. Five species of mustelids, Mustela
cf. erminea, Mustela vison, Mustela eversmanni
beringiae ssp. no\-., Gulo guJo, and Taxidea taxus,
are reported from late Pleistocene deposits near
Fairbanks, Alaska. This is the first record of the
steppe ferret in the New World. It is closely related
to, if not conspecific with, Mustela nigripes, the
black-footed ferret. The northernmost occurrence
of Taxidea taxus is reported. The woI\ erine, bad-
ger and ferret material is characterized by large
size, and some of tlie specimens are the largest
known for the species. The Fairbanks area was
never glaciated, and the grassy steppes of this
refiigium supported a large assemblage of Pleisto-
cene mammals.

INTRODUCTION

Remains of Pleistocene mammals are
abundant in the frozen sediments of central
Alaska, and at least 39 species are known.
Many species of carni\'ores were associated
with the large assemblage of herbivores that
inhabited the Alaskan refugium in the late
Pleistocene. Large carnivores, Arctodus si-
mtis, Ursus arctos. Panther a ho atrox, Homo-
fherium serum, and Canis lupus dominated
the scene, but the small carnivores — foxes,
dhole, lyn.x, and the mustelids — were an im-
portant part of the fauna. Five species of
mustelids, Mustela cf. erminea, Mustela
vison, Mustela eversmanni beringiae ssp.
nov., Gulo gulo, and Taxidea taxus are now
known from the Fairbanks area.

Fossil collecting began in the Fairbanks
area with the advent of gold mining in 1928.
In 1929, the University of Alaska, under the

^730 Magnolia St., Den\er, Colorado 80220.

presidency of C. E. Bunnell, initiated its
well known program of collecting the fossils
exposed during the mining operations. The
university had little money for such ven-
tures, but Childs Frick of the American
Museum of Natural History agreed to fi-
nance the program, and his support con-
tinued until the middle 1950's (except
during the war years when little mining was
done ) . Otto W. Geist was in charge of col-
lecting the fossils. Thousands of specimens
were collected, but unfortunately, because
of the methods of collection, stratigraphic
infonnation is almost entirely lacking. Since
the Universitv of Alaska had neither the
space nor the comparative material, almost
all of the specimens were shipped to the
Frick Laboratory at the American Museum
of Natural History. There, a few groups
were studied, but most of the material was
put in storage. The Mustelidae was one of
the neglected groups, and until 1973, when
Anderson reported the presence of ferret,
only badger and wolverine were recorded
in the faunal lists (Pewe, 1957).

The Fairbanks area, where the fossils were
collected, lies between 64°45' and 65° N lat-
itude, and is situated on the north side of
the broad Tanana River valley at the base of
the hills that make up part of the Yukon-
Tanana Ri\'er upland (see Fig. 1). Rising
380 to 545 meters above the nearly flat
floodplain of the Chena and Tanana rivers,
are the low rounded hills of the uplands.
Loess, derived from the floodplain and the
glacial outwash plains, covers the ridges

Bull. Mus. Comp. Zool., 148(1): 1-21, April, 1977 1

Bullcfiii Miiscrun of Comparative Zoology, Vol. 148, No. 1

I48°00

65*00

kilometsrs
— Moin Roads

Figure 1. Map of the Fairbanks area.

from a depth of about a meter on tlie sum-
mits to about 30 meters on the middle
slopes. The upland valleys are filled with
from three to 30 meters of gravel overlaid
by three to 90 meters of colluvial silt. The
floodplain is underlaid by several hundred
feet of interbedded lenses of silt, sand, and
gravel (Pevve, 1957). The Fairbanks area
was never glaciated, but glaciers from
the Alaska Range to the south came within
80 kilometers of the present cit\ of Fair-
banks.

The Quaternary in central Alaska is char-
acterized by alternating periods of deposi-
tion and erosion of gravel and silt, by
warming and cooling of the climate, and by
the formation and melting of the perma-
frost. Gold-bearing gravels were deposited
in the creek valleys early in the Quaternary,
and were later covered by loess and organic
debris which became perennially frozen.
Solifluction, the movement of moisture-sat-
urated soil downhill during periods of thaw-
ing, was a major factor in the entombment
of animal and plant remains. The fossil-
laden silt eventually came to rest in the

valleys, and was subsequently covered with
more loess, and the entire mass became fro-
zen. Today permafrost covers much of the
Fairbanks area. Needless to say, mining
and collecting fossils in this perennially
frozen muck was, and still is, difficult. The
fossils were exposed as the miners, using
hydraulic methods, removed the frozen
overburden from the gold-bearing gravels.

Since most of the bones were transported
before burial, mummies and complete skele-
tons are rare, and most of the specimens are
disarticulated. Although a few pre-Wiscon-
sinan deposits are known (Pewe and Hop-
kins, 1967), the majority of specimens are
late Wisconsinan in age.

The mustelid material is generally well
preserved, although some of the teeth are
broken. The bones vary in color from light
to dark brown, and there is no trace of the
blue mineral, vivianite, on any of the mate-
rial I examined. The specimens consist en-
tirely of skulls and mandibles. A femur of
Giilo was listed in the field notes, but the
specimen could not be found.

Pleistocene Mustelidae • Anderson

ACKNOWLEDGEMENTS

I wish to express my deep appreciation to
Dr. Richard H. Tedford for letting me study
the Alaskan mustelids in the Frick Col-
lection. Berv'l Ta>lor and George Krochak
of the Frick Laboratory, American Museum
of Natural History, assisted me in locating
specimens and field data. Russell D. Guth-
rie, University of Alaska, showed me some
of the collecting areas near Fairbanks, and
I would like to thank him and his wife for
their generous hospitality during my visit
to Fairbanks. For permission to study col-
lections in their care, I am indebted to John
A. \Miite, Idaho State University; Peter
Robinson, University of Colorado Museum;
Charles S. Churcher, University of Toronto;
and C. R. Harington, National Museum of
Canada. My sincere thanks go to Barbara
Lawrence and Charles Mack, Museum of
Comparative Zoology; John L. Paradiso and
Clyde Jones, Bureau of Sport Fisheries and
Wildlife, National Museum of Natural His-
tory; Richard G. Van Gelder, American
Museum of Natinal Histoiy; William H.
Burt, University of Colorado Museum; and
Robert S. Hoffmann, Museum of Natural
History, University of Kansas, for making
comparative material available to me. Bjorn
Kurten, University of Helsinki, permitted
me to use some of his raw data on Giilo
giilo. Erica Hansen, Idaho State University,
made the illustrations for Figures 1-3; Ms.
Dehlin, formerly of the Frick Laboratory,
executed Figures 4 and 5 for Childs Frick
some years ago. This research was sup-
ported by NSF Grant GB 31287 awarded to
Professor Bryan Patterson, Harvard Univer-
sity, and is part of a study of Pleistocene
mammals of North America.

ABBREVIATIONS

AMXH — American Museum of Natural Histoiy,
F:AM — Frick Collection, American Museum of

Natural History
ISUM — Idaho State University Museum
KU — Museum of Natural History, University of

Kansas
MCZ — Musemn of Comparative Zoolog\', Harvard

University
NMC — National Musemns of Canada

UA — Unixersity of Alaska

UCM — University of Colorado Museimi

USNM — National Museum of Natural History

I — incisor i

C — canine ' with superscript (upper) or

P — premolar ( subscript (lower) tooth

M — molar '

max. — maxillary

R — right

L— left

N — number in sample

O.R. — observed range

M — mean

S.D. — standard deviation

Mustela sp. cf. M. erminea Linnaeus
Short-tailed Weasel or Ermine
Figure 2 A

Material: Late Pleistocene, F:AM 49340 L ramus
w/C-M.; F:AM 49341 R ramus w/C-M.; F:AM
49348 R ramus w/C-Mi; F:AM 49349 frag. L
ramus w'Pa-i, Fairbanks area, Alaska.

Comparati\'e Material: Mustela erminea arctica,
Recent, Alaska AMNH 17939, 21917-19, 21921-
22, 31369, 31379. KU 2975-76. Mtisiela rixosa
eskimo Recent, Alaska AMNH 31383-84, 42811-
13, 42815-18. Northwest Territory AMNH
29212.

Four small weasel mandibles were found
in the collections from the Fairbanks area.
Guthrie (personal communication) believes
they were preserved in the nests of ground
squirrels, Spermophilus parryi. The coro-
noid process is missing in all of the speci-
mens. The teeth of three of the specimens
are slightly worn, but F:AM 49348 shows
moderately worn dentition. Comparison
with Recent specimens of Mustela erminea
arctica ( Merriam ) and Mustela rixosa es-
kimo (Stone), the two subspecies found in
central Alaska today, shows that the Pleisto-
cene specimens most closely resemble Mus-
tela erminea arctica. Table 1 shows that
measurements of tooth row length, length
of Ml, length of trigonid of M,, and width
of the talonid of Mi of die Pleistocene man-
dibles fall within the observed range of
Mustela erminea arctica and exceed the
observed range of Mustela rixosa eskimo.

In his monograph on American weasels,
Hall (1951) noted that the basilar length
of the skull of Mustela erminea measures

4 BtiJIcliii Museum of Comparative Zoology. Vol. 14S, Ko. 1

Table 1. Measurements, in mm, of Mustela

ERMINEA AND Mv STELA RIXOSA FROM ALASKA.

30cin

O.R.

Depth of ramus below P3-4

F:AM, Late Pleistocene

3.0-3.2

3.05

M. e. arcfica (Recent)

(

3.3-4.3

4.02

2.6-3.5

2.96

M. r. cskhno (Recent)

2..3-3.4

2.86

2.2-2.5

2.40

Depth of rannis below Mi

_.,

F:AM, Late Pleistocene

2.5-3.3

3.03

M. e. arctica (Recent)

3.5-1.8

4.40

2.9-3.7

3.26

M. r. cskimo (Recent)

2.7-3.5

3.13

2.5-2.9

2.72

Length C-M2

F:AM, Late Pleistocene

11.9-12.0

11.93

M. e. arctica ( Recent )

12.0-15.8

14.52

11.4-12.7

11.90

M. r. eskimo (Recent)

9.5-10.8

10.10

9.0-9.7

9.22

Length Mi

F:x'\M, Late Pleistocene

4.3-4.6

4.46

M. e. arctica (Recent)

4.5-5.6

5.01

4.1-4.7

4.33

M. r. cskimo (Recent)

3.4-3.8

3.55

3.1-3.5

3.25

Length Mi trigonid

F:AM, Late Pleistocene

3.0-3.3

3.20

M. e. arctica (Recent)

3.1-3.8

3.60

2.9-3.4

3.06

M. r. cskimo (Recent)

2.4-2.6

2.50

2.2-2.5

2.30

Width Ml talonid

F:AM, Late Pleistocene

1.2-1.4

1.33

M. e. arctica (Recent)

1.2-1.9

1.57

1.2-1.3

1.23

M. r. eskimo (Recent)

1.0-1.2

1.08

0.9-1.0

0.92

more than 32.5 mm in males and more than
31.0 mm in females; in Mu.sfeld rixosa the
basilar lengtli of the .skull is less than 32.5
mm in males and 31.0 mm in females. Un-
fortimately, he did not include any measure-
ments of weasel mandibles. Table 1 shows
that there is overlap in measurements be-
tween the two species, and this, coupled
with pronounced sexual dimoiphism and
geographic variation, can lead to uncer-

lOcm

Figure 2. A. Mustela of erminea (F:AM 49340); oc-
clusal and lateral views of mandible. B. Mustela vison
(F:AM 30821); occlusal and lateral views of mandible.

tainty in the identification of cranial mate-
rial.

The American ermine is considered to be
only subspecifically distinct from the Old
World animal. The taxonomic status of the
least weasel is uncertain. Some workers
(see Jones, 1964) regard Mustela rixosa as
only subspecifically distinct from the Eur-
asian Mustela nivalis. But, in Sweden, the
two species live side by side without inter-
breeding (Kurten, personal communica-
tion). Until detailed comparative and
statistical studies are done on both the Old
and New World populations, I am recogniz-
ing Mustela rixosa as a distinct species.

The ancestry of Mustela erminea can be
traced back to the late Pliocene in Europe,
and the species probably reached North
America in late Blancan or early Irving-
tonian times. The earliest known North
American occurrence is from the Cudahy

Pleistocene Mustelidae • Anderson

Table 2. Measurements, in mm, of Mustela
visoN FROM Alaska.

OR.

S.D.

Depth of ramus
below P3-4
F:AM 30821 1 7.8 — —

M. V. ingens
(Recent, Alaska) 16 6.1-8.3 7.45 ± .17 .71

Depth of ramus

below Ml -2
F:AM 30821 1 8.2 _ _

M. V. ingens

(Recent, Alaska) 16 7.0-9.4 8.26 ± .19 .79

Length Mi
F:AM 30821 1 7.8 — —

M. V. ingens
(Recent, Alaska) 16 7.4-9.0 8.14 ± .12 .49

Length Mi trigonid
F:AM 30821 1 5.3 — —

M. V. ingens
(Recent, Alaska) 16 5.1-6.1 5.75 ± .08 .32

Widtli Ml talonid
F:AM 30821 1 3.3 — —

M. V. ingens
(Recent, Alaska) 16 2.7-3.8 3.31 ± .08 .34

fauna, and ermines have been reported from
several late Pleistocene and postglacial lo-
calities.

Mustela vison (Schreber) Mink
Figure 2 B

Material: Late Pleistocene, F:AM 30821 frag-
ment of left ramus with P3-M1, Fairbanks Creek,
Alaska.

Comparative Material: Musiela vison ingens. Re-
cent, Alaska, MCZ 34165. USNM 6531-32,
7115, 8646, 8696-99, 8702-06, 8708-09, 14463,
20814. Yukon Territory MCZ 34517-18.

A single mandible of Mustela vison is
known from the Fairbanks area. The jaw is
broken off anterior to the second premolar,
and the coronoid process is eroded as is the
labial side of the condyle. The sharply
pointed cusps of the teeth show slight wear.
Each tooth is surrounded by a well devel-
oped cingnlum, and there is an incipient
metaconid on Mi. Table 2 shows that mea-

surements of the specimen fall within the
observed range of Mustela vison ingens
(Osgood), the extant subspecies found in
the area today. It is the largest .subspecies
of Mustela vison. No moiphological differ-
ences were observed between the specimen
and the comparative material.

The specimen can be distinguished easily
from the Mustela eversmanni mandibles by
the incipient metaconid and wider talonid
on Ml, and the longer and narrower P4.
Table 3 shows other differences separating
mink from ferret.

Although records of Mustela vison extend
back to the late Irvingtonian Cudahy fauna,
Meade County, Kansas (Getz, 1960), mink
are not common in Pleistocene deposits.
Since they are found only along streams and
lakes, the presence of mink in a fauna is a
good indicator of nearby permanent water.

Mustela (Putorius) eversmanni Lesson
Steppe Ferret

Material: Late Pleistocene, Fairbanks area, Alaska,
F:AM 49336 anterior half skull w/R P'^, L F,
C, P"", P* broken, Ester Creek. F:AM 49337 L
mandible W/ C-M^, Cripple Creek. F:AM 30827
frag. L ramus w/L, C, P., P.-Mi, Cripple Creek.
Mustela eversmanni Recent, MCZ 23705, 24737,
25333, 40939-40, 54604. USNM 22191, 188449,
259792. AMNH 57338, 60102, 85382. Mustela
eversmanni miehnoi USNM 38365, 172631,
175439, 175441. AMNH 45605-06, 84312.
Mustela putorius Recent, MCZ 3702, 24665,
24738 25352. USNM 792, 1851, 22394, 115213-
214 121248, 123629, 152668-670, 152673-676,
154158, 319222-223. AMNH 36631-32, 69520,
11962l', 163437.

Mustela nigripcs. Late Pleistocene, Litde Box
Elder Cave, Converse Count>', Wyoming, UCM
21916-18, 21922-24, 21950-52, 21957, 21959,
21962, 21965-70. 21972, 21975, 21977-78,
21980, 21983, 21985, 21989-90, 22010-11,
22022-23, 22151.

Mustela nigripes. Recent, MCZ 4184, 42723,
43727. KU 1487, 1593, 7146, 10177, 11077,
14411. AMNH 1203, 40078, 41994, 42567,
70590, 121610, 140397. UCM 59, 10658, 10660.
USNM 14580, 21066, 21965, 21976, 22311,
22427 22929, 30064-66, 32771, 34977, 35011,
35016-18, 35088, 35376. 65061, 83992-994,
110772 122620, 155475, 168744, 188450-453,
188455-458, 199737, 201945, 211513, 224450,
228233 228789, 232400, 234118, 234138,

6 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

Table 3. Comparison between Mustela eversmanni-nwripes and Mustela vison, cranial charac-
ters.

Variate

M. eversmanni-nigripes

M. vison

Palate

Basiocciput

Basicranium

Auditory bullae

Mastoid bullae

Auditory meatus

Infraorbital foramen

Canines, upper and lower

Mandible

Inferior margin of jaw at

angle
Premolars
Ml

Wide between canines

Narrow

Well-defined tube extend-
ing from foramen ovale
to anterior margin of au-
ditory bullae

More inflated

Inflated

External opening large

Small

Relati\ ely large

Short, broad

Relatively short protocone

Inner lobe not expanded

Relatively short and thick-
Broad, flattened

Relatively short, broad

Metaconid absent, talonid
narrow

Relativelv small

Narrow between canines

Wide

Area between foramen

ovale and auditory

bullae is flat

Less inflated

Not inflated

External opening small

Large

Relatively small

Long, narrow

Relatively long protocone

Inner lobe expanded

Relatively long, slender

Pointed, less flattened
Relatively long, narrow
Incipient metaconid,

talonid wide
Relatively large

134970-971, 234973, 241014, 243799, 243818-
820, 243909-910, 243990, 245641, 247073,
251453, 285877, 287321, 289498.

Anderson ( 1973) reported the presence of
ferret in central Alaska. Additional studies
show that the material is referable to Mus-
tela eversmanni, the steppe ferret, an animal
closely related to, if not conspecific with,
Mustela nigripes the black-footed ferret.
This is the first record of Mustela evers-
inanni in North America.

Mustela everstnanni beringiae^ ssp. nov.
Beringian Ferret
Figure 3

Type. F:AM 49336 anterior half of skull with
right F'-\ alveoli of P ^ C; left V, C-F % P^ bro-
ken, alveoh of I' ^ MS Ester Creek, T 1 S, R 2
W, about 16 km west of Fairbanks, Alaska 64°
50'N, 148°W. Fairbanks D-2, D-3 Quadrangles.
Collected in 1938.

Hypodigm. Type plus F:AM 49337 left mandible
with C-M2, alveoli of h-:>., Cripple Creek. F:AM

* beringiae — from Beringia, the enormous un-
glaciated land mass extending from westeiTi Alaska
to northeastern Siberia during the Pleistocene.

30827 fragment of left ramus with L, C, P., P4-
Mi, alveoli of I1-2, Ps, Cripple Creek, T 1 S, R 2

W, west of Fairbanks, Alaska.

Distribution. Known only from late Pleistocene de-
posits near Fairbanks.

Diagnosis. Large ferret; facial region broader than
Mustela eversmanni michnoi; massi\e postorbital
processes; pronounced postorbital constriction;
broad palate; tooth row crowded; enlarged ca-
nines.

A broad facial region characterizes the
skull, and measurements of the breadth
across the canines, carnassials, interorbital
region, and postorbital processes exceed
those of all the ferrets I have measured or
have seen referred to in the literature. The
skull belonged to an adult animal — the teeth
are moderately worn, the nasal and palatine
sutures are obliterated, and the sagittal
crest is well developed. The nasal opening
is large, and the opening of the small infra-
orbital foramen is an elongated oval. Ex-
tending from the tips of the broad postor-
bital processes are distinct ridges that unite
in the region of the postorbital constriction
to form the sagittal crest. The area between
the postorbital processes and the constric-

Pleistocene MusxELroAE • Anderson 7

lOcm

Figure 3. Mustela eversmanni beringiae ssp. nov.

A. (F:AM 49336 Type) occlusal view of partial skull.

B. (F:AM 49337) occlusal and lateral views of man-
dible.

tion is long and straight. The skull is bro-
ken just posterior to the constriction.

The upper teetli are crowded; there is no
diastema between the canine and P-, and
P^ is set obliquely in the jaw with the talon
slightly overlapping P^. The incisor row is
curved, and the width from the outer edge
of the alveolus of the right l^ to the outer
edge of the alveolus of tlie left P measures
8.9 mm; this compares with a mean of 6.79
mm (N 7, O.R. 5.8—7.6 imn) for Recent
Mustela eversmanni michnoi and 6.40 mm
(N 67, O.R. 5.4—7.2 mm) for Recent Mus-
tela nigripes. The canine is relatively long

and slender, and P^ is relatively short and
broad. Measurements of P^ fall within the
observed range of measurements taken on
the steppe ferret, and do not show any pro-
portional differences. Only the alveolus of
the left M^ is preserved; it shows that the
inner lobe of the tooth was narrow.

The left mandible, F:AM 49337, is per-
fectly preserved, only the incisors are miss-
ing (Fig 3B). The teeth are moderately
worn and are close together. Po is set
obliquely in the jaw and P4 slightly overlaps
Ml. As with the other species of ferrets, the
lower premolars are relatively short and
broad, Mi shows no trace of a metaconid,
the talonid of Mi is ridged and relatively
narrow, and Mo is small. F:AM 30827, a
partial left ramus broken off behind Mi, has
more heavily worn teeth than F:AM 49337.
The jaw is relatively massive in both speci-
mens, the length of the tooth row exceeds
those of Mustela eversmanni michnoi in my
sample, but measurements of the teeth fall
within the obser\'ed range of the other fer-
rets measured. ( See Table 4. )

Comparison of the Alaskan material with
both Pleistocene and Recent Mustela {Pu-
torius) nigripes Audubon and Bachman,
Recent Mustela ( Putorius ) eversmanni Les-
son and Recent Mustela (Putorius) putorius
Linnaeus showed that the specimens most
closely resemble Mustela (Putorius) evers-
manni michnoi Kashchenko, 1910, the South
Transbaikalian Siberian Polecat. This is the
largest subspecies and it inhabits the steppes
south and west of Lake Baikal and neigh-
boring areas of Mongolia ( Stroganov, 1962 ) ,
The skull of this subspecies shows a broad
facial region, pronounced postorbital con-
striction, and a crowded tooth row.

Stroganov (1962) reports that Mustela
eversnuinni shows more geographic varia-
tion than other ferrets, and about 20 sub-
species are recognized. For this reason, I
used only specimens labeled Mustela evers-
manni michnoi and Mustela eversmanni
larvatus, a synonym of the former (see
Ellerman and Morrison-Scott, 1966:265) in
my statistical analysis.

8 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

Table 4. Measurements, in mm, of Mustela eversmanni and Mustela nigripes.

O.R.

S.D.

Breadth across rostrum (C-C)

F:AM 49336

M. eversmanni niicluwi Recent

M. nigripes Recent

Little Box Elder Ca\ e Pleist.

Breadth across carnassials ( P'-P* )

F:AM 49336

Af. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Interorbital Ijreadth

F:AM 49336

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Breadth across postorbital processes

F:AM 49336

M. eversmanni michnoi Recent

M. nigripes Recent

Litde Box Elder Cave Pleist.

Breadth across postorbital constriction
F:AM 49336

M. eversmanni michnoi Recent
M. nigripes Recent

Litde Box Elder Cave Pleist.

Length C-M^

F:AM 49336

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Length F

F:AM 49336 ,

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Width P'

F:AM 49336

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Length P'

F:AM 49336
M. eversmanni michnoi
M. nigripes Recent
Little Box Elder Cave

Width P* protocone
F:AM 49336
M. eversmanni michnoi
M. nigripes Recent
Little Box Elder Cave

Recent
Pleist.

21.8

—

15.8-19.8

17.81 ± .53

1.40

15.1-19.6

16.80 ± .11

.91

16.8

—

28.4

21.0-26.2

24.57 ± .60

1.61

21.2-25.8

23.63 ± .12

1.02

21.8

16.4-19.3

17.64 ± .39

1.02

14.9-19.5

17.08 ± .11

1.05

18.0-21.4

19.70

—

26.3

20.0-23.6

21.84 ± .50

1.33

18.4-23.9

20.96 ± .16

1.42

21.8-25.4

23.60

14.9

10.2-15.9

12.42 ± .67

1.78

9.8-16.0

12.41 ± .12

1.09

12.5-16.3

13.96

22.1

18.7-22.1

21.00 ± .43

1.15

17.5-21.9

19.74 ± .29

2.62

ca21.4

4.4

3.8-4.6

4.28 ± .09

.26

3.5-4.2

3.86 ± .02

.17

3.8-4.4

4.1

2.6

2.0-2.7

2.31 ± .09

.25

1.9-2.5

2.18 ± .02

.16

2.2-2.3

2.25

8.5

7.0-8.5

8.10 ± .19

.52

6.7-8.0

7.41 ± .03

.28

7.3-8.1

7.71 ± .11

.30

4.4

3.4-4.3

3.80 ± .12

.31

3.2 4.0

3.67 ± .02

.18

3.5-4.1

3.77 ± .08

.21

Table 4. (contintied)

Pleistocene Mustelidae • Anderson 9

O.R.

S.D.

Length of mandible
F:AM 49337

M. eversmanni michnoi Recent
M. nigripes Recent

Little Box Elder Cave Pleist.

Height of mandible
F:AM 49337

M. eversmanni michnoi Recent
M. nigripes Recent

Little Box Elder Cave Pleist.

Depth of jaw below Ps-i
F:AM 49337, 30827
M. eversmanni michnoi Recent
M. nigripes Recent

Little Box Elder Cave Pleist.

Depth of jaw below Mi 2
F:AM 49337

M. eversmanni michnoi Recent
M. nigripes Recent

Little Box Elder Cave Pleist.

Length C-M2

F:AM 49337

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Lengtli of Ml

F:AM 49337, 30827

M. eversmanni michnoi Recent

M. nigripes Recent

Little Box Elder Cave Pleist.

Length of Mi trigonid
F:AM 49337, 30827
M. eversmanni michnoi Recent
M. 7iigripes Recent

Little Box Elder Ca\e Pleist.

Width Ml talonid
F:AM 49337, 30827
M. eversmanrn michnoi Recent
M. nigripes Recent

Little Box Elder Cave Pleist.

45.0

38.5 46.6

43.25 ± 1.51

4.00

36.4-45.6

42.14 ± 0.24

2.09

35.8-42.2

38.91 ± 1.48

3.64

22.0

20.0-23.1

21.41 ± .39

1.03

17.1-22.5

20.55 ± .13

1.17

18.4-21.8

19.75 ± .45

1.10

9.2

8.3-10.7

9.31 ± .21

.56

7.3-9.5

8.49 ± .07

.63

7.1-10.0

8.66 ± .21

.90

9.8

7.3-10.5

9.14 ± .23

.63

7.1-9.6

8.67 ± .07

.63

7.7-9.9

8.55 ± .16

.70

26.4

23.9-25.1

24.50

21.5-26.1

24.09 ± .11

1.03

21.5-25.3

23.37 ± .43

1.36

8.5-8.8

8.65

7.6-9.6

8.94 ± .25

.65

7.3-9.1

8.27 ± .04

.40

7.3-9.0

8.24 ± .09

.45

6.1-6.4

6.25

5.5-6.8

6.35 ± .18

.48

5.2-6.4

5.85 ± .02

.25

5.3-6.4

5.92 ± .19

.91

2.4-2.5

2.45

2.1-2.6

2.45 ± .08

.23

2.1-2.6

2.34 ± .01

.13

2.0-2.5

2.26 ± .02

.12

Extant steppe or Siberian ferrets are
found in steppe and forest-steppe zones of
Eurasia, from Hungaiy and Yugoslavia to
the Amur region of Siberia, south to the
plains of central Asia, Mongolia and north-
east China (Stroganov, 1962).

There is still disagreement as to the ge-
neric and specific status of Old World fer-

rets. Pocock (1936) and Ellerman and
Morrison-Scott (1966) recognize a single
species, MiisteJa (Putorius) piitoriiis. How-
ever, Russian scientists (Ognev, 1931 and
Stroganov, 1962), with larger samples to
work with, recognize Putorius putorius and
Putorius eversmanni as distinct species.
Stroganov lists the following cranial chai-ac-

10 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

ters as distinctive of Mustela eversmanni: a
larger, bulkier skull, appreciable constric-
tion of the postorbital region, and a longer
facial region. In addition, the canines and
carnassials are relatively larger than those
of Mustela puforius. There are also pro-
noiuiced differences in body size, coloration,
and habitat of the two species. Mustela pu-
forius inhabits forest biotopes and farm-
lands; Mustela eversmanni lives on the
steppes and seldom enters forests. In areas
where the ranges of the two species overlap,
the two forms remain distinct.

Although postorbital constriction is cor-
related with increasing age in most muste-
lids, skulls of Mustela putorius do not show
the pronounced consti'iction seen in the
other species. The mean of the measure-
ments of postorbital constriction of Mustela
putorius in my sample is 16.31 mm (N 24,
O.R. 12.6-18.2 mm); this compares with
a mean of 12.41 mm for Mustela eversmanni
and Mustela ni gripes (see Table 4).

Pocock (1936:715) noted "the close simi-
larity in all dimensions" of a male skull of
Mustela eversmanni from the Altai, and a
male skull of Mustela nigripes from Mon-
tana. I took 26 measurements on 19 skulls
of Mustela eversmanni and on 79 skulls of
Mustela nigripes; there were no significant
differences in size between the two species
(see Table 4). The only difference that
appeared on scatter diagrams was a nar-
rower basioccipital region in Mustela ni-
gripes. Both species inhabit steppe regions,
have a long sinuous body, and similar color-
ation. Musteln nigripes has never been
abundant on the Great Plains, and today it
is considered to be an endangered species.
Lhilike the steppe fen-et, which feeds on a
wide variety of small animals, the black-
footed ferret feeds primarily on Cynomtjs.
The geographic range of Cynomys and
Mustela nigripes are nearly identical and
the two species are associated in most
Pleistocene localities except Old Crow
River; Cynomys has not been reported from
Fairbanks.

The Pleistocene history of Mustela evers-

manni is poorly known, especially in Siberia.
It is reported from late Pleistocene deposits
in Europe; whether late middle Pleistocene
ferrets are Mustela putorius or Mustela
eversmanni is uncertain. Both species may
be derived from the smaller early middle
Pleistocene species, Mustela (Putorius) stro-
meri Konnos (Kurten, 1968).

The earliest record of Mustela nigripes is
from an upper Illinoian deposit in Clay
Countv, Nebraska, and it is known from
Sangamon deposits in Nebraska and at
Medicine Hat, Alberta. Wisconsinan rec-
ords include Old Crow River, Yukon Ter-
ritory; Orr Cave, Montana; Jaguar Cave,
Idaho; Little Box Elder Cave, Wyoming;
Chimney Rock, Colorado; Isleta Cave, New
Mexico; and Moore Pit, Texas. The speci-
men from Burnet Cave, New Mexico (see
Schultz and Howard, 1935) is a juvenile
with deciduous dentition; whether it is a
mink or a ferret cannot be determined. The
partial right ramus, NMC 16323, from Old
Crow River, Locality 65, may be referable
to Mustela eversmanni beringiae.

Ferrets entered the New World from Si-
beria, spread across Beringia, and then ad-
vanced southeastward to the Great Plains
through ice-free corridors. Kalela ( 1940, in
Kurten, 1957) reported that during the
period from 1880 to 1940, Mustela putorius
extended its range in Finland from the
Karelian Isthmus north to central Ostro-
bothnia and west to the Gvilf of Bothnia.
The rate of migration was 7.5 km annually
or 750 km in a century. When climatic
conditions permitted, this rate was probably
applicable for ferrets spreading across Si-
beria and into the New World.

The question of conspecificity between
Mustela eversmanni and Mustela nigripes
is yet to be resolved. That the two species
are closely related cannot be doubted, but
imtil detailed comparative and statistical
studies are made on the large collections of
Mustela eversmanni in Soviet institutions;
these data are compared with the informa-
tion already compiled on Mustela nigripes;
and behavioral and chromosomal studies

Pleistocene Mustelidae • Anderson 11

are undertaken on both species, I regard
them as distinct.

Gulo gulo (Linnaeus)
Figure 4

Wolverine

Material: Late Pleistocene, Fairbanks area,
Alaska, F:AM 30795 skull with complete denti-
tion, Goldstreain. F:AM 30796 anterior VL> skull
w/R I--^', P'-M\ L P, F -^ Ester Creek. F:AM
30797 L ramus w/C, Pj-Mi, top of gravel at 21
Coldstream, 40 feet below original surface. F:
AM 30798 skull and associated jaw sxmphvsis
w/R C, P^-M\ L P, P-C, P^; jaw symphysis
w/R and L C, P^Mi, Old Eva Creek. F:Ax\I
30799 partial anterior V2 skull w/R C, P'-M\
Cripple Creek. F : AM 30800 R max. w/P*, Ester
Creek. F:AM 30805 frag. R ramus W/P3-M1,
Fairbanks Creek. F:AM 30806 L ramus w/P..-
Mi, No. 2 Coldstream stripping area. F:AM
30807 frag. R ramus W/P3-M2, No. 2 Coldstream
stripping area. F:AM 30808 R ramus W/P^-Mi,
Cripple Creek. F:AM 30809 L ramus w/C, P3-
Mi, Cripple Creek. F:AM 30810 L ramus w/Po-
M2, Engineer Creek. F:AM 30811 frag. R ramus
W/P4-M1, Cripple Creek. F:AM 68003 R max
w/C broken, P^, M^ and assoc. frag. R. ramus
w/Mi-=, Gold HiU. F:AM 68005 frag. R ramus
W/P3-4, Gold Hill.

Comparative material: Postglacial, Moonshiner
Cave, Bingham County, Idalio, ISUM 19585-
19399, 19643, 19667, 17 skulls and skull frag-
ments; ISUM 19601-19639, 39 mandibles.
Recent, Alaska, MCZ 47398-99, 48566-68, 50528.
AMNH 137270. Yukon Territory MCZ 34516.
Northwest Territory AMNH 3448-49, 3450,
34506-09, 37432-33. Measurements of 24 male
and 13 female skulls from Alaska (data from
Bjbrn Kurten).

The outstanding feature of the wolverine
material from the Pleistocene of Alaska is
the large size of the specimens. Compari-
sons wnth. samples from postglacial Moon-
shiner Cave, Idaho, and the Recent of
Alaska and northern Canada show that the
Alaskan Pleistocene specimens exceed the
others in all measurements except the inner
lobe of M^ the depth of the jaw below P0-4,
and the length of the lower tooth row ( C-
M2) (see Table 5). No attempt was made
to sex the fossil material. If I had, the size
differences would have been even more
pronounced.

The well preserved skull, F:AM 30795
(see Fig. 4A-B) from Coldstream, is the

largest wolverine skull known to me. The
condylobasal length measures 172 mm. The
largest specimen in my sample has a condy-
lobasal length of 151 mm, and the largest
specimen in Ognev's sample from the
U.S.S.R. measured 157.8 mm (1935:587).
Hall and Kelson (1959) give an observed
range of 127-140 mm for basal length for
the extant animal in North America. Other
cranial measurements of F:AM 30795 are
equally large, especially the mastoid
breadth, breadth across the carnassials, and
the approximate zygomatic breadth. The
well developed sagittal crest projects above
the dorsal surface of the skull, but unfortu-
nately, the overhanging projection is broken
off at the occiput. The powerful mastoid
processes point obliquely forward and
downward. A partial skull, F:AM 30796,
and a right maxilla, F:AM 30800, represent
skulls nearly as large as F:AM 30795.

The dentition of F:AM 30795 is complete
and shows moderate wear. The length of
the upper tooth row ( C-M^ ) measures 62.8
mm compared with 53.0 mm for the largest
specimen from Moonshiner Cave, and 55.6
mm for the maximum length in my Recent
sample. Stroganov (1962:245) gives an ob-
served range of 49-60 mm for the length of
the upper tooth row for Siberian Gulo gulo.
The incisors of F:AM 30795 are all worn
down to the same level. The tips of both
canines were broken off during the life of
the animal, and the remaining portions of
the fangs are worn smooth. The massive
cheek teeth are crowded, but do not over-
lap, and the tooth row is dominated by the
enormous carnassial with its small talon.

F:AM 30797 (Fig. 4C), a complete left
mandible lacking only the incisors, first pre-
molar and last molar, is the largest lower
jaw from the Fairbanks region. Its total
length, measured from the s\anphysis at the
alveolus of Ii to the most distant edge of tlie
condyle, is 112.8 mm, a measurement
larger than an>' in the postglacial or Recent
sample. The teeth are only slightly worn,
and are close together with P2 sitting
slightly obliquely in the jaw. The posterior

12 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

Table 5. Measurements,

IN MM, OF GULO

GULO.

O.R.

S.D.

Condylobasal length

F:AM Collection

139.4-172.0

155.70

Moonshiner Cave,

Id.

134.0-145.0

140.66

G. giilo. Recent

140.0-151.0

146.05 ± .58

3.26

132.6-141.0

135.00 ± .65

2.61

Zygomatic breadth

F:AM Collection

100.0-119.2

109.6

Moonshiner Cave,

Id.

G. giilo, Recent

98.3-113.2

105.0 ± .57

3.04

92.5-100.0

95.97 ± .42

1.69

Breadth across rostnin

(C-C)

F:AM Collection

41.4-48.6

44.92

Moonshiner Cave,

Id.

36.1-42.3

38.92 ± .94

2.50

G. giilo. Recent

40.0-43.9

42.51 ± .40

1.20

37.0-39.9

37.94 + .35

.99

Breadtli across carnass

ials (F-F)

F:AM Collection

67.1-76.7

72.40

—

Moonshiner Cave,

Id.

51.4-63.0

59.82 ± .73

2.07

G. gulo. Recent

63.6-69.3

66.97 ± .59

1.77

59.7-63.8

61.47 ± .49

1.40

Interorbital breadth

F:AM Collection

41.0 46.7

44.70

—

Moonshiner Cave,

Id.

36.7-44.6

40.60

G. gulo. Recent

39.1-45.4

41.38 ± .28

1.56

36.0-40.9

37.75 ± .15

.69

Breadth across postorbital processes

F:AM Collection

48.2-56.8

52.50

Moonshiner Cave,

Id.

44.0-53.5

48.37

G. gulo. Recent

45.7-54.3

48.61 ± .94

2.83

42.4-49.5

45.50 ± 1.05

2.79

Mastoid breadth

F:AM Collection

85.0-108.0

96.50

Moonshiner Cave,

Id.

76.2-87.3

81.18

—

G. gulo. Recent

83.1-94.6

90.05 ± 1.14

3.43

78.4-85.6

82.50 ± .82

2.19

Length C-M^

F:AM Collection

51.3-62.8

57.72

Moonshiner Cave,

Id.

46.4-53.0

50.47 ± .69

2.29

G. gulo. Recent

51.0-55.6

53.28 ± .42

1.28

43.4-^1.3

48.63 ± 1.43

4.05

Length P

F:AM Collection

21.6-23.4

22.70 ± .51

1.27

Moonshiner Cave,

Id.

18.0-22.3

20.36 ± .28

1.06

G. gulo. Recent

20.2-23.2

21.30 ± .13

.70

18.6-20.1

19.37 ± .09

.40

Widtli F protocone

F:AM Collection

12.6-13.5

12.96 ± .44

1.10

Moonshiner Cave,

Id.

10.4-13.4

11.58 ± .22

.86

G. gulo. Recent

11.4-13.1

12.31 ± .08

.47

10.6-12.1

11.15 ± .10

.43

Table 5. (continued)

Pleistocene Mustelidae • A^nderson 13

O.R.

S.D.

Width M^

F:AM Collecdon
Moonshiner Cave, Id.
G. gulo. Recent

Lengtli M^ constriction
F:AM Collection
Moonshiner Cave, Id.
G. gulo, Recent

Length M^ inner lobe
F:AM Collection
Moonshiner Cave, Id.
G. gulo. Recent

Length mandible

F:AM Collection
Moonshiner Cave, Id.
G. gulo. Recent

Depth of jaw below Ps-t
F:AM Collection
Moonshiner Cave, Id.
G. gulo. Recent

Depth of jaw below M1-2
F:AM Collection
Moonshiner Cave, Id.
G. gulo. Recent

Length C-M2

F:AM Collection
Moonshiner Cave, Id.
G. gulo, Recent

Length Mi

F:AM Collection
Moonshiner Cave, Id.
G. gulo, Recent

Length Mi trigonid
F:AM Collection
Moonshiner Cave, Id.
G. gulo. Recent

Width Ml talonid
F:AM Collection
Moonshiner Cave, Id.
G. gulo, Recent

13.5-15.8

14.80

12.0-14.2

13.39 ± .16

.72

13.7-15.7

14.45 ± .09

.51

12.5-13.9

13.05 ± .09

.40

6.3-6.6

6.46

5.1-6.2

5.72 ± .06

.29

5.6-6.5

5.99 ± .04

.26

4.9-5.9

5.53 ± .06

.29

7.1-9.0

8.06

6.6-8.5

7.56 ± .12

.51

7.3-9.7

8.20 ± .06

.34

6.8-8.1

7.22 ± .08

.35

96.0-112.8

105.82

89.2-107.0

95.41 ± .82

3.28

99.5-107.2

103.76 ± 1.28

3.85

94.2-99.4

95.86 ± .66

1.89

18.4-22.2

20.62 ± .44

1.33

16.0-21.4

18.64 ± .32

1.67

19.3-22.6

20.82 ± .94

2.83

17.9-19.0

18.42 ± .42

1.19

22.0-29.6

25.70 ± .55

1.66

20.0-26.1

22.21 ± .32

1.67

23.2-25.7

24.52 ± .58

1.76

19.6-22.3

21.35 ± .55

1.56

59.7-70.1

63.14 ± .74

1.97

54.7-64.6

58.82 ± .20

1.02

61.3-66.4

64.70 ± .54

1.64

57.5-62.1

59.86 ± .55

1.56

22.0-24.6

23.15 ± .57

1.81

18.5-23.1

20.80 ± .09

1.29

21.6-25.2

22.80 ± .32

1.74

19.5-22.0

20.85 ± .48

2.05

16.9-19.3

18.15 ± .27

.83

14.2-17.6

15.87 ± .15

.97

16.2-19.0

17.93 ± .28

.80

15.6-17.1

16.43 ± .17

.49

7.0-8.1

7.62 ± .13

.41

5.8-8.4

6.83 ± .29

.55

7.1-7.9

7.40 ± .09

.27

6.0-7.3

6.70 ± .18

.52

14 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

Figure 4. Gulo gulo (F:AM 30795) A. lateral and B. ventral views of skull; C. {F:AM 30797) lateral view of
mandible. Scale 1/1.

part of P4 is expanded. Mi is a massive
tooth watli a powerful trigonid and reduced
talonid; there is no trace of a metaconid.
F:AM 30797 was also found at Coldstream,
but it did not belong to the same individual
as F:AM 30795. Except for larger size, the
mandibles from the Pleistocene of Alaska
do not differ from the extant Gulo living in
the area today.

Kurten and Rausch (1959) in their study
of Alaskan and Fennoscandian wolverines
noted that a significant difference was

foimd between the two populations in the
length of NP measiu-ed at the constriction.
They found that the Recent specimens from
Alaska had a more strongly constricted M^
than those from Scandinavia. This is not
the case with the late Pleistocene Alaskan
specimens — the NP shows less constriction
than those from Scandinavia. One of their
fossil specimens from Europe also showed
this reduced constriction of M^ On a scat-
tergram the specimens from Moonshiner
Cave show nearly the same proportions as

Pleistocene Mustelidae • Anderson 15

the sample from Fennoscandia. Compari-
sons with other late Pleistocene samples of
Gulo are now being made.

Circumboreal in distribution, wolverines
inhabit tundra and taiga regions, and today
in America are found primarily in Alaska
and northern Canada. Wolverines are rare
in Pleistocene deposits. The earliest Amer-
ican records are late Irvingtonian from Port
Kennedy Cave, Pennsylvania and Cumber-
land Cave, Maryland. Wisconsinan deposits
containing Gulo include Old Crow River,
Yukon Territorv; Little Box Elder Cave,
Wyoming; Chimney Rock Animal Trap,
Colorado; Jaguar Cave, Idaho; and Fair-
banks. Wolverines show a gradual increase
in size during Rancholabrean times; post-
glacial and extant animals are smaller.

Gulo is descended from Plesiogulo, a
large Pliocene form with a less specialized
dentition that inhabited Eurasia and North
America. Gulo makes its first appearance in
early middle Pleistocene deposits in Europe
as a slightly smaller form called Gulo schlos-
seri Kormos. It gave rise to Gtilo gulo
which appears during the Mindel glaciation
in Europe and China. Gulo probably
reached America in the Kansan. American
wolverines were formerly considered to be
a distinct species, Gulo luscus (Linnaeus);
Kurten and Rausch ( 1959) showed that the
American population is only subspecifically
distinct from the Eurasian.

Taxidea taxus (Schreber) Badger
Figure 5

Material: Late Pleistocene, Fairbanks area, Alaska.
F:AM 30786 skull and associated mandible
\v/R r -, C-NP, L complete upper dentition,
R Ii 3, C, Pa-M,, L Ii-3, 1/2 P2, P3-M2, Gold-
stream. F:AM 30787 skull w, L C, P-M\ Gold-
stream. F:x\M 30788 R ramus \v,C, P4-M1, head
of Goldstream. F:AM 30789 L ramus w/Mi bro-
ken, Cleary. F:AM 30790 frag. L jaw, tooth-
less, Goldstream. F:AM 30826 L ramus w/Mi-2,
Cripple Creek. F:AM 30827 R ramus w/Pa-Mi,
Ester Creek. F:AM 30828 R ramus vv/P^, hU,
frag. Ml, Cripple Creek. F:AM 30829 L ramus,
toothless. Ester Creek. F:AM 30830 frag. R
ramus w/Po 4, Mi broken. Cripple Creek. F:AM
30831 L ramus w/C, Cripple Creek. F:AM

30832 L ramus w/C, P^Mi, all broken, Cripple
Creek. F:AM 30833 L max. w/F-M\ Cripple
Creek. F:AM 30834 L max. w/P'^, Cripple
Creek. F:AM 30835 anterior half skull w/R C,
P", L yoP^ Lower Goldstream. F:AM 30836
anterior half skull w/R C, F-M\ L VsF, Cripple
Creek. F:AM 30837 skull w/R P^ C-M\ L
I-" -^ C, 1/2 F, F, Ester Creek. F:AM 30837 A
frag. R ramus W/P2-M0, Ester Creek. F:AM
30838 partial skull w/R and L F-M\ Ester
Creek. F:AM 30839 L ramus w/C, P3-M1, Fair-
banks Creek. F:AM 30840 frag. R ramus w/Mi,
Cripple Creek. Field numbers: F:AM 4493 L
max. w/F-M\ Gold Hill. F:AM 4717 R max.
w/C, F broken, F^*, Gold Hill. F:AM 4737 L
ramus W/P2-3, P4-M1 broken. Gold Hill. F:AM
6135 L ramus w/Mi 2, C-P4 broken off at roots.
Engineer Creek. F:AM 6411 jaw symphysis w/
R C-M2, L P3-4, all broken. Cripple Creek. F:
AM 68004 frag. R ramus w/P4, Gold Hill. U.A.
ace. no. 552 (on loan to Frick Laboratoi-v) skull
w/R and L F-M\ Cripple Creek.
Late Pleistocene, Little Box Elder Cave, Con-
verse County, Wyoming, UCM 21928. Postgla-
cial, Moonshiner Cave, Bingham Count\', Idaho,
ISUM 19650, 19671-79, 19682-85,' 19687,
19701-04, 19705, (36 specimens), 19706 (31
specimens), 19731-32, 19735-49, 19752, 19761-
64, 19766, 19769, 19771, 19773-75, 19777,
19780-81, 19795, 19799-19806, 19814-834.
Recent. Taxidea taxus ieffersonii MCZ 8517,
9223, 12402, 41389-90. UCM 5150, 5237, 5284,
5882, 6678, 10682-84, 10687. E.R. Warren col-
lection, not cataloged 2635, 9135. Taxidea taxus
herlandieii UCM 11548-550. UCM 3698, no
data.

Badgers are not found in Alaska today.
Their closest occurrence is along the Peace
River, lat. 58 °N, in northern Alberta
(Preble, 1908), about 1800 km southeast of
the Fairbanks area. During the late Pleisto-
cene, badgers inhabited the unglaciated,
grassy steppes of central Alaska and north-
ern Yukon (Gold Run Creek, Harington,
1970, and Dominion Creek, Harington, per-
sonal communication ) . Remains of Taxidea
outnumber the other Alaskan mustelids in
the Frick collection.

The Alaskan badgers are characterized by
large size. The condylobasal length of U.A.
ace. no. 552 is 144.6 mm, a measurement
that exceeds all other Pleistocene, postgla-
cial, and Recent records. The condylobasal
length of die large skull from Little Box
Elder Cave, UCM 21928, (see Anderson,

16 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

Table 6. MEAstTREMENTS, in mm, of Taxidea taxus

O.R.

S.D.

Condylobasal length

F:AM Collection

137.7-144.6

140.60

—

Moonshiner Cave, Id.

118.0-129.2

124.55 ± .61

2.04

Taxidea taxus. Recent

114.0-132.0

122.47 ± 1.22

4.89

Zygomatic breadth

F:AM Collection

90.6-100.1

95.35

—

Moonshiner Cave, Id.

72.4-80.1

76.96 ±1 1.31

3.95

Taxidea taxus. Recent

72.4-87.7

77.91 ±1.18

4.74

Breadth across rostrum (C-C)

F:AM Collection

38.4-46.2

41.96 ± 1.20

2.94

Moonshiner Cave, Id.

32.1-37.8

35.00 ± .36

1.53

Taxidea taxus. Recent

30.5-37.9

34.15 ± .46

2.06

Breadtli across carnassials (P'-P*)

F:AM Collection

45.2-49.6

47.40

Moonshiner Cave, Id.

38.5-44.6

40.94 ± .41

1.74

Taxidea taxus, Recent

37.7-44.4

40.85 ± .38

1.68

Interorbital breadth

F:AM Collection

32.9-39.7

36.91 ± 1.07

2.61

Moonshiner Cave, Id.

25.6-35.0

29.90 ± .53

2.47

Taxidea taxus. Recent

24.6-31.4

27.50 ± .37

1.64

Breadth across postorbital processes

F:AM Collection

39.9-44.6

42.28 ± .65

1.60

Moonshiner Cave, Id.

31.8-37.0

35.03 ± .28

1.32

Taxidea taxus. Recent

30.8-40.3

35.25 ± .53

2.40

Mastoid breadth

F:AM Collection

90.0-91.4

90.80

Moonshiner Cave, Id.

66.0-83.3

73.89 ± .64

2.22

Taxidea taxus. Recent

70.2-86.8

76.39 ± 1.09

4.64

Length C-M^

F:AM Collection

40.7-47.3

44.74 ± .68

2.04

Moonshiner Cave, Id.

36.1-44.3

40.32 ± .33

1.93

Taxidea taxus. Recent

35.3-43.0

39.87 ± .38

1.73

Length P^

F:AM Collecti(m

7.3-8.6

7.87 ± .13

.39

Moonshiner Cave, Id.

6.5-7.3

6.90

Taxidea taxus, Recent

6.2-7.5

6.64 ± .10

.40

Length F

F:AM Collection

11.5-13.9

12.88 ± .24

.75

Moonshiner Cave, Id.

10.8-13.5

11.86 ± .08

.53

Taxidea taxus. Recent

10.3-13.5

11.73 ± .18

.80

Widtli P' protocone

F:AM Collection

10.2-12.1

11.15 ± .20

.63

Moonshiner Cave, Id.

9.0-11.9

9.91 ± .09

.57

Taxidea taxus. Recent

9.0-11.3

10.05 ± .14

.61

Width M^

F:AM Collection

10.1-12.0

10.93 ± .19

.56

Moonshiner Cave, Id.

9.1-11.8

10.07 ± .09

.63

Taxidea taxus. Recent

9.3-11.6

10.28 ± .15

.67

Table 6. (coNTiNtrEo)

Pleistocene Mustelidae • Anderson

O.R.

S.D.

Length NP inner

F:AM Collection
Moonshiner Cave, Id.
Taxidca taxus. Recent

Length mandible

F:AM Collection
Moonshiner Cave, Id.
Taxidca taxus. Recent

Depth of jaw below P3-4
F:AM Collection
Moonshiner Cave, Id.
Taxidea iaxtis, Recent

Depth of jaw below M1-2

F:AM Collection
Moonshiner Cave, Id.
Taxidea taxus. Recent

Thickness of jaw below Mi
F:AM Collection
Moonshiner Cave, Id.
Taxidea taxus. Recent

Length C-M2

F:AM Collection
Moonshiner Cave, Id.
Taxidca taxus. Recent

Length P*

F:AM Collection
Moonshiner Cave, Id.
Taxidea taxus, Recent

Length Mi

F:AM Collection
Moonshiner Cave, Id.
Taxidea taxus. Recent

Length Mi trigonid
F:AM Collection
Moonshiner Cave, Id.
Taxidca taxus, Recent

Width Ml talonid

F:AM Collection
Moonshiner Cave, Id.
Taxidea taxus. Recent

9.5-12.6

10.95 ± .39

1.09

9.7-12.5

10.90 ± .11

.70

9.6-12.9

11.16 ± .21

.93

95.2-109.0

99.01 ± 1.70

4.81

73.7-98..3

85.63 ± .65

4.56

78.8-93.5

86.18 ± .99

4.32

15.1-21.3

17.85 ± .41

1.73

12.0-18.1

14.42 ± .13

1.08

12.0-16.3

14.35 ± .27

1.23

19.5-26.0

22.58 ± .37

1.56

16.2-21.5

18.54 ± .14

1.18

15.(^22.0

18.28 ± .35

1.58

8.1-12.0

10.23 ± .27

1.14

7.0-9.4

8.18 ± .13

.71

6.5-8.6

7.65 ± .18

.71

51.2-59.3

54.64 ± .71

2.15

43.4-53.6

49.14 ± .33

2.33

44.6-52.0

48.79 ± .48

2.09

8.2-9.8

9.14 ± .19

.57

7.2-8.6

7.88 ± .20

.53

7.3-8.7

8.15 ± .12

.48

13.1-15.2

14.,38 ± .21

.73

11.6-14.9

13.20 ± .20

.89

12.,3-14.6

13.55 ± .17

.66

7.7-10.5

9.48 ± .37

.99

7.7-9.9

8.70 ± .15

.66

8.3-9.9

8.87 ± .11

.45

5.6-7.2

6.34 ± .14

.47

4.9-6.5

5.80 ± .09

.43

5.3-7.0

6.03 ± .12

.51

196S) measures 142.2 mm. Table 6 shows
that the largest specimen from Moonshiner
Cave has a condylobasal length of 129.2
mm, and the largest Recent skull in my
sample measures 132.0 mm. Long (1972)
gives an observed range of 121. .5-139.9 mm
for the greatest length of the skull of Tax-

idca taxus jeffersonii, the largest extant sub-
species. Other big late Pleistocene badgers
are known from Dominion Creek, Yukon
Territory, Rancho La Brea, McKittrick, and
Maricopa, California. Burnet Ca\'e and San-
dia Cave, New Mexico (personal observa-
tions ) . I am presently reviewing all of the

iJLtL

..ill Museum of Comparative Zoology, Vol. 148, No. 1

Figure 5. Taxidea taxus (F:AM 30786) A. lateral and B. ventral views of skull; C. occlusal view of upper
dentition; D. lateral view of mandible; E. occlusal view of lower dentition. Scale 1/1.

iMc'i.stoccne Taxidea material. Preliminary
studies indicate that all of the material is
referable to Taxidea taxiis, but subspecific
designation of the Blancan and late Rancho-
labrean material seems warranted. A trend
in the evolution of Taxidea during the
Pleistocene is a gradual increase in size
culminating in the huge Rancholabrean
forms; there was a slight decrease in size
during postglacial times, and this was fol-
lowed by a slight increase in size in Recent
times. The largest extant badgers are found
in the northern parts of their range.

All of the measurements taken on the
Alaskan material, except the length of M^
which is quite variable, exceed those in my
postglacial and Recent sample. Pronounced
size differences are noticed in zygomatic
breadth, mastoid breadth, and length of
mandible ( see Table 6 ) .

The well preserved skull, U.A. ace. no.
552, belonged to an adult animal. The low
broad skull is characterized by strong zygo-
matic arches, well developed sagittal and
lambdoidal crests, a wide occiput with
highly inflated tympanic bullae, and sepa-

Pleistocene Mustelidae • Anderson 19

rate paraoccipital processes. The incisors,
canines, and P^'s are missing; P^^-M^ are
moderately worn and close together. Three
other complete skulls, three partial skulls,
and four maxillary fragments are known
from the Fairbanks area. Large size is char-
acteristic of all of them. F:AM 30837 and
30787 have condylobasal lengths of 139.5
mm and 137.7 mm respectively; F:AM
30836 has a rostmm breadth of 46.2 mm
compared to 43.6 mm for U.A. ace. no. 552.

Eighteen badger mandibles were found
in the Fairbanks area. Of these, F:AM
30832 is the largest, the total length of this
massive jaw measures 109 mm; this com-
pares with a measurement of 98.3 mm for
the largest specimen from Moonshiner Cave
and 93.5 mm in my Recent sample. The
teeth of F:AM 30832 are heavily worn, and
this plus the great size indicate advanced
age. The teeth of several of the specimens
are broken. Moi-phologically, the specimens
do not differ from the Recent sample. As
Hall ( 1944 ) noted, the number of accessory
cusps on the talonid of Mi is extremely
variable in Recent badgers; this is also true
in the Alaskan population.

Geographic variability, sexual dimor-
phism, and individual variation are pro-
nounced in badgers. Most fossorial of the
Mustelidae, badgers inhabit plains and open
forests where friable soil is available for
digging. Their diet consists of insects and
small vertebrates, especially rodents. Al-
though badgers are inactive during cold
spells, they are not true hibernators. The
presence of badgers in Alaska during the
late Pleistocene indicates a milder climate
then, for todav their northern distribution
is limited by subarctic conditions. Hall
(1944) cites the vicissitudes of the boreal
climate as the major factor preventing inter-
continental exchange of Old and New World
badgers, and he postulated that if this ex-
change had occurred, the genus Meles
would be found in North America as well
as Eurasia, and Taxidea would be restricted
to the southern latitudes of the New World.
At the time Hall wrote this (1944), the

Alaskan badgers were unknown. Why they
did not spread farther West across Beringia
is unknown.

Badgers are common in Pleistocene de-
posits in western United States, and a few
have been recovered from sites in the East
including Cumberland Cave, Maryland;
Welsh Cave, Kentucky; Baker Bluff, Ten-
nessee; and Peccaiy Cave, Arkansas. The
probable ancestor of Taxidea is PUotaxidea
nevadensis ( Butterf ield ) known from Hem-
pillian faunas in Nevada and Oregon. It
was smaller and had larger tympanic bullae
than Taxidea. Today Taxidea taxus is found
from southern Canada to southern Mexico
and from the Pacific Coast east to Michigan
and Ohio.

CONCLUSIONS

During the late Pleistocene at least five
species of mustelids inhabited an ice-free
refugium in interior Alaska. Although strati-
graphic information is lacking, all of the
mustelid material is believed to be Wiscon-
sinan in age. Pewe and Hopkins (1967) do
not list any species of mustelids from pre-
Wisconsinan age deposits in the Fairbanks
region, and carbon-14 dates obtained on
bison, musk ox, and mammoth material from
the same area fall between 12,460 and
>40,000 years B.P. (B. Taylor, personal
communication ) .

The mammalian fauna of Alaska and
northeastern Siberia was similar during the
Wisconsinan, since biogeographically, it
was one vast area. At the height of the
glaciation, many species of animals ranged
across the Beringian refugium unable to
move onward because of the ice. Some of
them, for example. Saiga, Bos (yak). Tax-
idea, and Megalomjx, did not extend their
range, but many others, mainly the Eurasian
immigrants, moved southward when the
ice-free corridors were open. Hopkins
(1967) postulated that an ice-free corridor
probably existed in the Yukon Territory,
northern British Columbia, and northern
Alberta during the mid-Wisconsinan, a pe-
riod of mild chmatic conditions between

20 Bulletin Museum of Comparative Zoology, Vol. 148, No. 1

35,000 and 25,000 years ago; the corridor
was closed from about 22,000 years to at
least 14,000 years ago; and then it reopened
again after the Bering land bridge had been
drowned by rising sea levels. Thus, move-
ments of animals to and from the Beringian
refugium took place in mid-Wisconsinan
and very late Wisconsinan/postglacial
times.

As Hopkins (1967) and Guthrie (1968)
postulated, grasslands must have been more
extensive in the refugium during the late
Pleistocene in order to have supported the
enormous numbers of herbivores that lived
there. The remains of thi'ce obligatory graz-
ers. Bison, Eqiius, and Mammiithus, make
up more than 85 per cent of the fossils col-
lected in the Fairbanks area, and the pres-
ence of many plains dwellers including
Taxidea taxus and Mtistela eversmanni fur-
tlier supports this hypothesis.

Large size was characteristic of many
species during the Pleistocene, and remains
of Gido gido and Toxidea toxus from the
Fairbanks deposits are the largest recorded.
This may be an example of Bergmann's
principle — that the same species of warm-
blooded animal tends to be larger in the
colder parts of its range — but an abundant
food supply and few enemies may also have
been factors.

The extinction or extirpation of many
members of the Beringian fauna about
10,000 years ago was probably due to mul-
tiple factors including abrupt changes in the
climate which resulted in changes in the
vegetation (for example, an increase in the
tundra-taiga and bogs at the expense of
grasslands). This affected the large mam-
mals more than it did the small ones. Of the
mustelids, Taxidea taxus and Mtistela evers-
manni disappeared from Alaska, but sur-
vived in areas much farther south; Gtdo
gtdo, Miistela vison and Musteki erminea
still inhabit the area today. Man was un-
doubtedly a factor in the extinction of some
species, but it is doubtful that he had any-
thing to do with the disappearance of two
of the Alaskan mustelids.

REFERENCES

Anderson, E. 1968. Fauna of the Little Box
Elder Cave, Converse County, Wyoming. The
Camivora. Univ. Colorado Stud., Eartli Sci.
No. 6: 1-59.

. 1973. Ferret from the Pleistocene of

central Alaska. Jour. Mammal. 54(3): 778-
779.

Ellerman, J. R. AND T. S. C. Morrison-Scott,
1966. Checklist of Palearctic and Indian
mammals 1758-1942. 2nd ed. London, Brit.
Mus. (Nat. Hist.).

Getz, L. 1960. Middle Pleistocene carnivores
from southwestern Kansas. Jour. Mammal.
41: 361-365.

Guthrie, R. D. 1968. Paleoecology of the large
mammal community in interior Alaska during
the late Pleistocene. Amer. Midi. Nat. 79(2):
346-363.

Hall, E. R. 1944. A new genus of American
Pliocene badger with remarks on the relation-
ships of badgers of the Northern Hemisphere.
Carnegie Inst. Washington, Publ. 551: 9-23.

. 1951. American weasels. Univ. Kansas

Publ. Mus. Nat. Hist. 4: 1-466.

and K. Kelson. 1959. Mammals of

North America. Vol. II. New York, Ronald
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Harington, C. R. 1970. Ice Age mammal re-
search in the Yukon Territory and Alaska. In
Early Man and environments in northwest
North America. Student Press, Univ. Calgary.
pp. 35-51.

Hopkins, D. M. 1967. The Cenozoic history of
Beringia — a syntliesis. In Hopkins, D. M.
(Ed.), The Bering Land Bridge, Palo Alto,
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Jones, J. K. 1964. Distribution and taxonomy of
mammals in Nebraska. Uni\ . Kansas Publ.,
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KuRTEN, B. 1957. Mammal migrations, Ceno-
zoic stratigraphy, and the age of Peking Man
and the australopitliecines. Jour. Paleontol.
31(1): 215-227.

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London, Weidenfeld and Nicolson. 317 pp.
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parisons between North American and Euro-
pean mauunals. Acta Arctica. 11: 1-44.

Long, C. A. 1972. Taxonomic revision of the
North American badger, Taxidea faxiis. Jour.
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Ognev, S. I. 1931. Mammals of easteiTi Europe
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. 1935. Mammals of U.S.S.R. and adja-
cent countries. (Israeli Program Scientific
Translation, 3: 72-90-1- Table 24. 1962).

Pewe, T. L. 1957. Permafrost and its effect on

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life in the North. 18th Biology Colloquium,
Corvallis, Oregon, pp. 12-25.

-, AND D. M. Hopkins. 1967. Mammal re-
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Hopkins, D. M. (Ed.), The Bering Land
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ScHULTZ, C. B. AND E. B. HowARD. 1935. The
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us ISSN 0027-4100

BuiLetln OF THE

Museum of

Comparative

Zoology

Studies on the Deep Sea Protobranchia

(Bivalvia); The Family Tindariidae and

the Genus Pseudotindaria

H. L. SANDERS AND J. A. ALLEN

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 2
14 APRIL 1977

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chusetts, 02138, U.S.A.

STUDIES ON THE DEEP SEA PROTOBRANCHIA (BIVALVIA);'
THE FAMILY TINDARIIDAE AND THE GENUS
PSEUDOTINDARIA

H. L. SANDERS- AND J. A. ALLEN^

Abstract. In the present paper we have erected
a new family of Protobranchia, the Tindariidae, to
inckide those nuculanoid bivalves that lack a si-
phon but bear papillae on their posterior margin
edge, ha\'e a single loop of tlie hind gut that pene-
trates the right side of the mantle, and possess a
palp with few ridges. A new genus, Pseiidoiindaria,
is created for those nuculanoid protobranch bi-
valves that have shell morphologies essentially
similar to the Tindariidae but with soft parts that
are markedly different. Siphons are present, the
hind gut is a complex configuration of loops and
coils on either side of the body and it does not
penetrate the mantle, and palp ridges are numer-
ous. The morphologies of the hard and soft part
anatomies, the horizontal and vertical distributions,
size-frequency histograms, and reproductive pat-
terns of the tindariid and pseudotindariid species
in our Atlantic samples are discussed. Two new
species are described. On tlie basis of shell mor-
phology, Tindaria and Pseudotindaria can be
interpreted as recent descendants of the Paleozoic
ctenodont Protobranchia.

INTRODUCTION

Tlie objects and aims of our researches on
the fauna of the deep sea, and on the Proto-

^ This research was supported by grants GB 563,
GA 31105 and GB 36554 from the National Science
Foundation, GR 3, 812 from the Natural En\iron-
ment Research Council, and from the Royal Society
of London. Contribution no. 2983 from the Woods
Hole Oceanographic Institution.

- Woods Hole Oceanographic Institution, Woods
Hole, Massachusetts, U.S.A.

^ Dove Marine Laboratory, University of New-
castle upon Tyne, Cullercoats, England.

branchia in particular, have been given in
the prologue to these studies, (Sanders &
Allen, 1973). This is the third paper in a
series, all of which illustrate the initial
problems that had to be resolved in our
analysis and reappraisal of the deep-sea
protobranch bivalves of the Atlantic.

x\s noted in the first paper (Sanders &
Allen, 1973), tlie tindariid protobranchs
represent an anomalous group within the
Order Nuculanoidea, having features that
divide them sharply from the remainder of
the order. Verrill & Bush (1897), Theile
(1935), Voices (1967) and Knudsen (1970)
all place the genus Tindaria {= Tyndaria)
in the family Malletiidae, even though Ver-
rill & Bush' (1898) stated that "the genus
Tindaria differs so widely from MoUetia
and other genera that it seemed necessary
to establish a new subfamily (Tindarinae)
for it."

Dall (1895) included the following sub-
genera: Tindaria, Tindariopsis, Neilonella
and Vseudoglomus in the Malletiidae and
Knudsen (1970) retained the genus Neilon-
ella there as well. Theile (1935) divided
the genus Tindaria into t\vo sections each
with a single subgenus, Tindaria and
Pseudoglomus.

Because Tindaria differs so markedly in
its morphology from both Pseudoglomus
and Neilonella as well as other members
of the family Malletiidae, we propose that
a new family be erected, the Tindariidae.

Bull. Mus. Comp. Zool., 148(2): 23-59, April, 1977 23

24 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

The genera Fseudo glomus, Neilonella and
Malletia will be the subjects of future
papers.

The present study shows that some spe-
cies, although having typical tindariid shells,
have greatly different anatomies from the
true tindariids. On the basis of these pro-
found anatomical differences we find it
impossible to include these species within
the same family. We propose that those
forms having papillae around the incurrent
aperture, lacking well defined siphons, and
having a hind gut configuration consisting
of a single deep loop on the right side of the
body, be included in the family Tindariidae.
Excluded from the family are tliose forms
with a well-developed siphon and complex
hind gut configuration that extends on both
sides of the body. These we include within
a new genus P sen dotind aria, described here.
The precise affinities of this genus will be
deferred to a futvue paper. This interpreta-
tion in no way conflicts with earlier defini-
tions of the genus Tindaria, all of which
mention posterior papillae (e.g. Theile,
1935).

TINDARIIDAE New Family

The family is characterized as follows:
valves rounded, ovate, robust, swollen,
somewhat unequilateral, concentrically
lined; umbo medially and somewhat ante-
riorly directed; hinge line strong, supporting
a series of well-developed teeth that are
continuous beneath the umbo; ligament
external and opisthodetic; posterior sensory
tentacle present or absent; true siphons
lacking; incurrent region of mantle edge
fringed with elongate papillae; palps small;
sorting ridges of the palp broad, few in
number; palp proboscides large, elongate;
gill axis somewhat oblique to the antero-
posterior axis of the body; gill filaments
few; hind gut, with lumen of large diameter
and a single typhlosole, making a single
loop to the right side of the body and pene-

trating into the mantle to a greater or lesser
extent; 'byssal' gland small.

Although they show several features that
sharply differentiate them from other proto-
branchs, the tindariids clearly fall within i
our definition of the Order Nuculanoidea
(Sanders & Allen, 1973). For example, all \
species of the family have a posterior in-
current current and an anterior mantle sense

organ.

Tindaria Bellardi 1875

Type species T. arata Bellardi, by monotypy

Tindaria is the sole genus of the family,
the generic characters of the genus are
those that define the family. Many species
are listed in the literature ( see Smith, 1885;
Clarke, 1962; Knudsen, 1970) but, for rea-
sons that will become obvious, unless the
soft parts have been described many of
these species cannot be placed in the genus
with confidence. Of the features that dis-
tinguish both the genus and family, the
most distinctive are 1) the lack of siphons
and the long fringing papillae of the in-
current region, 2) the small size of the palps
relative to the size of the animal, 3) the
very few ridges on the inner surface of the
palps, 4) the oblique placement of the gill
in relation to the anterior-posterior axis, 5)
the relatively small number of gill filaments,
6) the small size of the 'byssal' gland, 7)
the single loop of the hind gut on the right
side of the body.

Tindaria callistiformis Verriil & Bush,
1897
Figures 1-11 & 27

Tindaria callistifoiDiis, Verriil & Bush, 1897. Anier.
I. Sci., p. 59, figs. 10, 20, 21 (Type locality: U.S.
Fish Conim. Sta. 2566; Lat. 37°23'N, Long. 63°
8'W, type specimen: U.S. Nat. Mus.); Verriil &
Bush, 1898. Proc. U.S. Nat. Mus., No. 1139, p.
881, pi. 78, fig. 1; 80, figs. 6, 7.

Previous records. Depth range = 3342 to 4795 ni.

North America Basin — 2 stations. Refs. Verriil &
Bush, 1897, 1898.

Present records. Depth range = 3305 to 5042 m.

Deep Sea Protobranchs • Sanders 6- Alien 25

Cruise

Station

No.

Depth

(m)

No. of
specimens

Latitude

Longitude

Gear

Date

Vorth America Basin

Chain 50

3806

622

38°0.7'N

69°16.0'W

30.6.65

Chain 50

3828

181

38°0.8'N

69°18.7'W

30.6.65

Chain 50

3832

882

37°59.2'N

69°26.2'W

5.7.65

Atlantis II

175

4667

36°36.0'N-
36°36.0'N

68 ° 29.0' W-
68°31.0'W

29.11.67

Chain 50

4749

36°24.4'N

67°56.0'W

4.7.65

Atlantis II

121

4800

35°50.0'N

65°11.0'W

21.8.65

Atlantis II

122

4833

35°50.0'N-

35°52.0'N

64°57.5'W-
64°58.0'W

21.8.65

Atlantis II

123

4853

37°29.0'N

64°14.0'W

22.8.65

Atlantis II

124

4862

37°26.0'N-

37°25.0'N

63°59.5'W-
63°58.0'W

22.8.65

Chain 50

5042

Angola

34°41.0'N

Basin

66°28.0'W

2.7.65

Atlantis II

197

4592-4597

10°29.0'S

9=04.0'E

21.5.68

Atlantis II

196

4612 4630

Argentine

10°29.0'S

Basin

9°04.0'E

21.5.68

Atlantis II

259A

3305-3317

37°13.3'S

54M5.0'W

26.3.71

Atlantis II

256

3906-3917

Guiana

37M0.9'S

Basin

52°19.5'W

24.3.71

Knorr 25

307

3835-3862

12°35.4'N
12°40.8'N

58°59.3'W
59°09.2'W

3.3.72

Knorr 25

288

4417-4429

11°02.2'N
11°03.8'N

55°05.5'W
55°04.8'W

25.2.72

Knorr 25

287

4934-4980

13°16.0'N

13°15.8'N

54°52.2'W
54°53.1'W

24.2.72

Specific description. We can make but
few additions to the excellent description
of shell morphology given by Verrill & Bush
(1898). The small medial teeth immedi-
ately below the umbo insert in a dorsal arc,
away from the ventral edge of tlie hinge
plate (Figs. 1 & 27a). The external liga-

Iment extends posteriorly in the mid-line of
. the escutcheon to about the posterior limit
of the umbo at the insertion of the ninth
tooth of the posterior plate series. The an-
terior ligament is short, not extending be-
)ond the beak of the umbo.
I The incurrent region has four or five
' papillae on each side and lacks mantle
fusion between the posterior apertures. The
gills are small, with 10 to 14 gill plates
on each demibranch, and 7 to 10 palp

ridges. The hind gut passes close to the
anterior adductor muscle, but is not con-
tiguous with it. The anterior adductor mus-
cle is slightly larger than the obliquely
orientated posterior adductor muscle (Fig.

2).

Morphology of the soft ports. No de-
tailed account of tlie soft part anatomy has
been given hitherto. Lack of mantle fusion
and siphons (note, siphons may be formed
in the Nuculanoidea without fusion of the
mantle tissues, Yonge, 1959) is reminiscent
of the condition in tlie Nuculoidea ( Sanders
and Allen, 1973). However, unlike tlie
members of that order, the posterior mantle
edge is highly specialized and divided into
excurrent, incurrent and feeding regions in
addition to the ventral pedal gape (Fig. 3).

26 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

digestive gland

oesophagus

Figure 1. Tindaria callistiformis Verrill & Bush. In-
ternal views of the left and right valves.

The exciirrent region occupies a very small
portion of the posterior mantle margin. A
narrow channel is defined by the ventral
edge of the posterior adductor muscle and
by a pair of low ridges, formed by the inner
muscular mantle folds, to which the attenu-
ate distal end of the gill axes ai-e attached.
The anus is positioned opposite this narrow
channel. There is no development of the
inner muscular fold of the mantle edge to
form an incomplete siphon as Knudsen
(1970) described for some other parts. The
incurrent region is wider than the excurrent
and defined by four to six pairs of short
conical papillae, the number depending on
the size of the specimen. On the right side
immediately adjacent to the lower papilla
there is a single sensory tentacle. Tlie

Figure 2. Tindaria callistiformis Verrill & Bush.
Semidiagrammatic drawing of the body and mantle
organs as seen from right and left sides.

papillae are developed from the middle sen-
sory lobe. In cross section, the papillae are
radially divided into 12 to 15 haemocoelic
cavities running the length of the papillae
with longitudinal muscle fibres at the
center.

Ventral to the incurrent region there is a
feeding aperture which is foraied by the
extended overlapping and folded portions
of the inner and middle mantle folds, and
it is through this aperture that the palp ii
proboscides are extended (Fig. 3). The
inner muscular fold in the region of the \i
posterior apertures is much broader than ^
elsewhere. Gland cells are present in the .
outer mantle epithelium in the region im-
mediately posterior to the feeding aper-
ture. These extend anteriorly, although less
densely, to the inside of the muscular fold
in the region of the pedal aperture ( Fig. 4 ) . '

Deep Sea Protobranchs • Sanders 6 Allen 27

faecal rod

tentacle

folded
manfle

Figure 3. Tindaria callistiformis Verrill & Bush. Detail of the posterior mantle edge spread open and viewed
from the ventral side; the limits of the various apertures indicated wWh dashed lines.

An anterior sense organ, derived from the
middle sensory fold, is well-developed. The
left sense organ is somewhat larger than the
right, the latter positioned immediately be-
low the \entral limit of the hind gut loop.
For a comparatively robust shell, the adduc-
tor muscles are small and characteristically
situated close to the pallial line and very
near the shell margin. The 'quick' and
'catch' parts of the adductor muscles are
clearly defined. The anterior muscle is
circular in outline, while the posterior ad-
ductor is oval with the long axis oblique to
the antero-posterior shell axis.

The gill axis lies parallel to the posterior
dorsal margin of the shell and thus some-
what obliquely to the anterior-posterior axis
of the body and shell. The nrnnber of gill
plates is small, varying with the size of the
animal. They are widely separate and alter-
nate on either side of the axis ( Fig. 2 ) . The
gill extends across the posterior third of the
body to the ventral side of the excurrent
region. Posteriorly, the gill plates do not
extend much beyond the posterior limit of
the body, thus the gill axes are extended

posteriorly. The anterior limit of the gill
axis is far removed from the insertion of the
palp. The gill axis is highly muscular, with
fibres extending its length and also reach-
ing \'ertically to each gill plate (Fig. 5). The
gill plates are finger-shaped and sub equal.
There is no fusion between mantle and gill,
and connections between the inner fila-
ments of the two gills are apparently lack-

ing-

The palps are remarkable for their small
size and the small number of ridges ( seven
to nine ) , the exact number being dependent
on the size of the animal ( Fig. 6 ) . The palp
ridges are broad, high and deeply grooved
on the mid-anterior face. The most poste-
rior ridge is well anterior to the posterior
thickened edge of the palp.

The foot is typically nuculanoid with a
well-defined neck at its junction with the
body. Within the neck are large pedal gan-
glia and associated statocysts. Small papil-
lae fringe the entire edge of the divided
sole. The heel is small and triangular and
internally there is a small 'byssal' gland with
paired apertiues opening at the junction be-

28 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Figure 4. Tindaria callistiformis \/e-ri\\ & Bush. Trans-
verse section through the mantle edge.

tvveen heel and foot. Numerous subepithe-
lial mucous glands open on to the sole of the
foot on either side and in the mid line ( Fig.

7).

The gut also is basically nuculanoid with
a single loop of the hind gut on the right
side of the body. The mouth is posterior
to and some distance from the anterior
adductor muscle. This may be due in part
to its displacement by the loop of the hind
gut, which passes close to the adductor mus-
cle. It may also have functional significance
in relation to the posterior ingress of food
material into the mantle cavity. The
oesophagus is long, first taking an ante-
rior course to the posterior dorsal edge of
the anterior adductor muscle where it is
displaced slightly to the left of the sagittal
plane. At this point it turns dorsally and
posteriorly to open on the left anterior side
of a huge stomach. The oesophagus is in-
flated close to its junction with the stomach
and the stomach occupies much of the body
space (Figs. 2 & 8). Although the stomach
is large, there are only six very low crested

outer g
plate

inner gill
plate

Figure 5. Tindaria callistlformis Verrill & Bush. Trans-
verse section through the left gill showing axial mus-
cles and outline of gill plates.

sorting ridges on its right side. Much of the
remainder of the stomach is lined with a
gastric shield which has a well-defined
tooth on the anterior dorsal side close to the
apertures of tlie three ducts of the digestive
diverticula. The combined mid gut and
style sac penetrate the neck of the foot, pass
tf) the posterior and left side of the pedal

F gure 6. Tindaria callistiformis Verrill & Bush. Lat-
eral view of inner surface of a proximal palp to show
detail of ridging.

Deep Sea Protobranchs • Sanders 6- Mlsn

£9

muscles

Figure 7. Tindaria callistiformis Verrill & Bush. Trans-
verse section of the foot to show position of the mus-
cles and pedal glands.

ganglion and join with the hind gut just

ventral to the latter. The liind gut turns

and follows a dorsal course parallel with

the style sac, with a shallow, ill-defined U-

bend posterior to the stomach. The hind

gut then forms a single loop to the right

I and the loop togetlier with some body tissue

i penetrates the right mantle to a position

I close to the palHal line. A typhlosole is

present throughout the length of the hind

gut.

The ganglia and their connectives are
large, as the visceral and cerebral ganglia
are elongated. A pair of statocysts dorsal
and postero-lateral to the pedal gangUa are
filled with small crystals (which are not
calcium carbonate ) lacking ducts to the out-
side. Transverse muscle fibres in the foot
are found dorsal and ventral to the ganglia
and the statocysts.

The kidney is small and multilobed.
Sexes are separate.

Size, Reproduction and Age. All three
stations from which large numbers of T.
calUstifonnis were collected showed similar
population histograms. Each was strongly
skewed to the left (Fig. 9). Yet, we might
not adequately be sampling the smallest
juvenile stages which could pass through
the 0.42 mm openings in our screens. The
eggs on hatching are at least 0.15 mm long
and the size of the metamoi-phosed post-

hind

gut

stomach

digestive
gland

tooth

of gastric

shield

digestive
duct

hind gut

Figure 8. Tindaria callistiformis Verrill & Bush. Trans-
verse section through the body to show detail of
stomach, hind gut and digestive gland.

lar\'ae settling onto the bottom (assuming
a lecithotrophic mode of reproduction)
must be somewhat larger. We feel our his-
tograms do not significantly distort the
length-frequency composition of T. coUisti-
formis at the sampling sites. Analysis of 60
specimens from Station 77, representative of
the size range, indicates that gonadal devel-
opment does not occur until this species
reaches a size of between 4.0 and 4.5 mm
total length. This was confirmed by exam-
ination of the specimens from Stas. 78 and
85 which show that all specimens larger
than 4.5 mm had some gonadal develop-
ment and the larger the specimen the more
mature was the gonad. Only in an excep-
tionally small percentage (2.6 to 3.0 per
cent depending on tlie sample ) of the total
population was any sign of gametogenesis
evident. Great disparit>' exists in the sex
ratio with only one female to eveiy five
males. The most mature female (6.5 mm

30 BiiUetin Museum of Comparative Zoology, Vol. 148, No. 2

total length) was dissected and the eggs
counted; these numbered 230 having a max-
imum length of 145 /x. From our past
obser\'ations on a wide range of abyssal
protobranchs, we would estimate this spec-
imen to be 3 4 mature. The eggs were
maturing simultaneously. There is no evi-
dence of brooding.

Growth rate measurements using --^Ra
chronology (Turekian et al., 1975) show
that Tindaria callistiformis having a length
of 4 mm are about 50 to 60 years old, while
the largest specimen, having a length of 8.4
mm, has an age of 100 years or longer.

The moderate degree of variation in shell
shape appears to have no intimate relation
to the size of the specimen. Thus, the
height/total length ratio varies from 0.62 to
0.83 over much of the size range (Figs. 10

and 11). However, if the five largest spec-
imens are ignored, there is a tendency for
the height total length ratio to decrease
with increasing size. Similarly, there ap-
pears to be no increase in length posterior
to the umbo with increasing length. Al-
though anterior in position, there is consid-
erable variation in the position of the umbo,
the extreme limits being between 55 per
cent and 76 per cent of the total length.
The maximum total length recorded is 8.4
mm.

Tindaria liessieri, new species
Figures 12-16 & 27

Holotype: MCZ 279902, from Atlantis 11, Cruise

31, Station 141, in 2031 m.
13t'pth range = 1739 to between 2051 and 2357 ni.

Station

Depth

No. of

C raise No

No.

(m)

Specimens

Latitude

Longitude

Gear

Date

W. Europe

Basin

Sar.sia

S-44

1739

Cape Verde

43°40.8'N

Basin

3°35.2'W

16.7.67

Atlantis II

138

1944-1976

10°36.0'N

17°52.0'W

4.2.67

Atlantis II

141

2131

10°30.0'N

17°51.5'W

5.2.67

Atlantis II

139

2099-2187

10°33.0'X

17° 53.0' W

4.2.67

Atlantis II

145

2105-2192

10°36.0'N

17°49.0'W

6.2.67

Atlantis II

144

2051-2357

10°36.0'N

17°49.0'W

5.2.67

Specific description. Shell stout, robust,
with strong, uniform, concentric ridges,
somewhat oval in outline and extended
posteriorly; uml^os anterior in position,
moderately swollen l)caks prominent and
strongly curved medio-anteriorly; escutch-
eon present and forms moderate concavity
( Fig. 27B ) ; elongate, narrow, external liga-
ment extends posteriorly along the escutch-
eon to about the insertion of the eleventh
tooth on the posterior hinge plate and
anteriorly to about the insertion of the
sixth or seventh tooth of the anterior hinge
plate; dorsal shell margin strongly convex;
antero-dorsal margin short, sloping rapidly
to form a continuous curve with the ante-
rior margin; postero-dorsal margin long,
sloping more gradually to form a smooth

curve with posterior margin; posterior end
narrow, e\'enly rounded; anterior end broad,
roimded, but with dorso-anterior shorter
than ventro-anterior margin; ventral margin
long and only slightly convex. Hinge plate
broad and strong with a continuous row of
teeth; posterior hinge plate long with about
21 to 22 teeth, 14 through 21 or 22 large and
robust, more medial teeth progressively
smaller and very reduced in size at conflu-
ence of the anterior and posterior hinge
plates; anterior hinge plate short and thick
l:)earing about 10 teeth, distal five large and
strong, more proximal teeth as on posterior
hinge plate become gradually smaller with
minute proximal teeth inserting dorsally
(Fig. 12).
Tindaria hessleri differs from T. callisti-

Deep Sea Protobranchs • Sanders 6- Allen 31

STA 78

175 SPECIMENS

i ^

5 6

STA 77
600 SPECIMENS

-r-

SHELL LENGTH (mm)

Figure 9. Tindaria callistiformis Verrill & Bush. Size frequency histograms of samples from two stations.
The dashed lines indicate the sizes at which gametogenesis is evident.

R 85-

■

■
■ ■

■

■
■

■

■ ■

■

m
m

S 65-

■ .

* ■

■

55-

i^i

^ 80-

|70-

oo°

° o o

|60-

°8 0

o o

o
o

§ 50-

1 1

3 4

/■ 1

SHELL LENGTH (mm)

Figure 10. Tindaria callistiformis Verrill & Bush. Graph showing height/total length (■) and umbo to poste-
rior margin/total length (O) plotted against total length.

32 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Figure 11. Tindaria callistiformis Verrill & Bush.
Growth series in lateral view.

formis in the following ways: T. hessleri is
less smoothly convex and more triangular in
outline; the ventral margin of the valve is
deepest directly beneath the umbo rather
than more posteriorly; the dorsal margin is
more strongly convex; the anterior margin
is not as broadly rounded; and the hinge
plates are less massive.

Incurrent region, three papillae on each
side; adductor muscles oval and equal in
size; gill small with 7 to 14 plates on each
side of the axis; palp with 8 to 10 ridges;
hind cfut extends into the mantle of the
right side to a position short of the pallial
line, part of hind gut lies adjacent to the
posterior face of the anterior adductor
muscle; visceral ganglion placed anterior
to the posterior adductor muscle.

Morphology of the soft parts. The
moiphology of T. hessleri is very similar to
that of T. callistiformis (Fig. 13). How-
ever, unlike the latter species, there is a

Figure 12. Tindaria hessleri Sanders & Allen,
nal views of the left and right valves.

Inter-

permanent excurrent aperture formed by
the fusion of the muscular lobe and the
inner part of the sensory fold of the mantle,
between the incurrent and excurrent regions
(Fig. 13). There are only three pairs of
papillae on each side of the incurrent region
and sectioned material suggests that mantle
fusion dorsal to this region involves the
homologue of a pair of papillae that have
fused together. There is no single sensory
tentacle but mantle extensions at the dorsal
side of the feeding aperture may possibly
serve the same function. The latter aper-
ture is well-developed with the muscular
and sensory folds of the mantle showing
considerable hypertrophy. These are folded
in preserved specimens but in life they
must be capable of considerable extension.

adductor

muscle

Deep Sea Protobranchs • Sanders 6- Alhr.

digestive gland

stomach

oesophagus

visceral ganglion

posterior

adductor

muscle

papillae

anterior mantle
sense organ

foot

^Ccc:.,

Figure 13. Tindaria hessleri Sanders & Allen. Semidiagrammatic drawings of the body and mantle organs
as seen from the left and right sides.

34 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

hindgut

axial
muscle

posterior

adductor

muscle

Figure 14. Tindaria tiessleri Sanders & Allen. Trans-
verse section through the posterior part of the body
and gills to show axial and pedal retractor muscles.

There is no mantle fusion either dorsal or
ventral to the feeding aperture, but aposi-
tion or overlapping of the mantle edges at
these points makes an efficient functional
separation. Adjacent to this region is a well-
defined area of acidophilic mucus secreting
cells at the surface of the inner mantle
epithelium. The adductor muscles are oval
in cross section with the longitudinal axis
vertical in the anterior muscle and oblique
in the posterior muscle. To the inside of the
inner muscular lobe and at 90 degrees to it
are a series of fine pallial retractor muscles
extending inwards for a short distance;
these are present along the entire perimeter
of the mantle (Fig. 13).

In specimens of a similar size there are
more gill filaments than in T. callisfiformis
and, as in the latter species, they are ar-
ranged alternately along the axis. Behind
the body, the tips of the filaments of the
inner demibranch are extended and fuse
with the filaments of the inner demibranch
of the opposite gill and with adjacent fila-
ments of the same demibranch, thus form-
ing a membranous junction. The filaments

Figure 15. Tindaria hessleri Sanders & Allen. Trans-
verse section through the right margin of the hypo-
branchial cavity showing the distribution of basiphilic
gland cells.

of the outer demibranches are also extended
and make a strong ciliary junction with the
mantle (Fig. 14). The gill axis is very mus-
cular and it must be concluded that in life
the gills form a pumping system in many
ways analogous to that of the Septibranchia.
The hypobranchial cavity is thus entirely
separate from the rest of the mantle, the
lateral mantle walls of the cavity being
lined with basiphilic gland cells (Fig. 15).
No fecal material passes into the mantle
cavity. When the gill is contracted the vol-
ume of the hypobranchial cavity is very
small.

The palps are very similar to those of
T. calUstiformis except that, depending on
the size of the animal, the number of ridges
is somewhat greater, and these lie relatively
closer together than in the latter species.
The palps ( and mouth ) lie far posterior to
the anterior adductor muscle. The course of
the gut is essentially the same as in the pre-
vious species but the lumen is relatively
smaller and does not extend as far into
the mantle, never reaching the inner mus-
cular lobe of the mantle — even in the
smallest specimens. The foot is similar to
that of T. calUstiformis, however, the 'bys-
sal' gland is somewhat larger. Nervous

Deep Sea Protobranchs • Sanders h- Mler. 3

system and kidney are as in the previous
species; sexes are separate.

Size and Reproduction. The small num-
ber of specimens and the opaqueness of the
shell makes an analysis of reproducti\e
potential similar to that given for T. callisti-
f or mis impossible. However, sections show
that a specimen of 2.5 mm total length
contains approximately 350 ova with a max-
imum length of 110 /jl. The ova are matiu-
ing simultaneously.

Relatively few shells (eight) were avail-
able for studies on dimensional variation
(Fig. 16). It appears to be of the same
order as Tindaria callistiformis and Pseiido-
tindaria galatheae with the height length
ratio varying from 0.69-0.81 and with
the total length/umbo to posterior margin
length varying from 58 per cent to 67 per
cent, the umbo being anterior in position.

It gi\'es us great pleasure to name this
species after Dr. R. R. Hessler, of the
Scripps Institution of Oceanography — col-

league, collaborator and friend — ^who has
contributed so significantly to our knowl-
edge of the abyssal fauna.

Tindaria cytherea (Dal I, 1881)
Figures 17-22

Nucida cytherea Dall, 1881, Bull. M.C.Z., 9, No.

2: 123 {Blake Station witliout number Yucatan

Strait, 640 fms. (=1171 m)). Holotype.

U. S. Nat. Mus. 63137.
Mallctia veneriformis Smith, 1885, Chall. Rep. 13:

246, pi. 20, figs. 9, 9a. ( Challenger Station 33,

off Bennuda, 435 fms ( = 796 m).)
Malletia cytherea (Dall), 1886, Bull. M.C.Z, 12,

No. 6: 254, pi. 8, figs. 1, la. [in part, figure

only.]
Mallctia amabilis Dall, 1889, Bull. M.C.Z., 18: 438,

pi. 40, fig. 8. [New name for Malletia cytherea

Dall 1886 in part, description only, not figure.]
Tindaria cytherea (Dall) Verrill & Bush, 1898,

Proc. U.S.N.M. No. 1139.
Previous records. Depth range = 714 to 1325 m.
North America Basin — 1 station. Ref. Smith, 1885.
Gulf of Mexico Basin— 1 station. Ref. Dall, 1886.
Caribbean Basin — 1 stations. Refs. Dall, 1881,

1886; SmiUi, 1885.
Present record. Depth 1000 m.

Cruise

Station

No.

Depth

(m)

No. of
Specimens Latitude

Longitude

Gear

Date

Panulirus

Bermuda #1

1000

North America Basin

1 32°16.5'N

66°42.5'VV

13.4.60

We have collected a single large speci-
men of a tindariid species off Bermuda in
1000 meters that is similar to or identical
with a number of forms described from the
same general region and depth. These are
T. cytherea, (Dall) T. anudnUs (Dall) and
T. veneriformis (Smith) (Fig. 17). Dall
(1886) synonymized T. cytherea and T.
veneriformis and later Verrill and Bush
(1898) synonymized T. cytherea and T.
amabilis, although Dall (1889) believed
that they were separate. Dall ( 1881, 1886,
1889) described two specimens, one in 1881
as Nuciila cytherea, and the second in 1886
as Malletia cytherea. In 1889 he stated that
the second specimen described in 1886 was
sufficiently different from the specimen
described in 1881 to warrant the erection of
a second species which he named M. ama-

l)His. The position is confused by the fact
that the specimen described as N. cytJierea
(Dall, 1881) is figured in his second paper
(Dall, 1886) while the specimen described
in 1886 is figtued in his third paper (Dall,
1889). We agree with Verrill and Bush
(1898) that these all refer to the same spe-
cies as indicated in the synonymy.

We have but a single specimen, measur-
ing 9.2 mm total length, height 7.6 mm.
Its description is as follows:

Description of Shell Shell stout, robust,
straw-colored, with strong, uniform, con-
centric ridges; oblong oval in outline with
anterior end slightly truncate, inequilateral;
umbo anterior in position, prominent, with
beaks strongly curved antero-medially; es-
cutcheon present; external hgament elon-
gate, moderately large, extending posteriorly

36 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Figure 16. Tindaria hesslerl Sanders & Allen. Growth
series in lateral view.

to about the insertion of the fourteenth
tooth of the posterior hinge plate series and
anteriorly to about the sixth tooth of the
anterior series; inner layer forms a small
crescent-shaped section immediately below
the beaks; dorsal shell margin strongly con-
vex with both antero- and postero-dorsal
shell margins steeply sloping away from the
umbos; long postero-dorsal margin forms a
smooth curve with the rather broadly
rounded posterior end; antero-dorsal mar-
gin forms a smooth curve with the
slightly truncated anterior end; ventral mar-
gin smooth, long, and only shallowly con-
vex; hinge plate strong, moderately broad
with teeth continuous beneath the umbo;
long posterior hinge plate bearing about 24
teeth which are directed primarily medi-
ally, proximal teeth smallest, distal teeth
increasingly larger to tooth 16 to 18, the
latter being large, long, and robust, there-
after teeth diminish in length but remain
robust; anterior hinge plate with about 14
medially directed teeth, teeth 6 to 13 strong
and long, more proximal teeth progressively
smaller.

Figure 17. Tindaria cytherea (Dall). A & B, internal
views of the valve (Type specimen USNM No. 63137,
Blake Collection, Yucatan Strait 640 fm); C, external
lateral view of specimen collected by authors.

Remarks. This specimen differs from the
descriptions given by Dall (1881, 1886) and
Smith (1885) primarily by the somewhat
more obvious external ligament and the
straight ventro-medial edge of the anterior
margin. In this respect it resembles the

Deep Sea Protobranchs • Sanders 6- Allen 37

^^^IIP"

Figure 18. Tindaria cytherea (Dall). Detail of the
hinge and teeth of the specimen collected by the
authors. A, left valve, umbonal region; semidiagram-
matic presentation of right valve in dorsal (B) and
posterior view/ (C).

t\^e specimen of M. amabilis (Dall) rather
than the type specimen of M. cytherea
(Dall) (Fig. 17) in which the ventro-me-
dial edge is curved. Shell proportion, shape
and thickness alter with growth in many
deep-sea protobranch species. These
changes are most evident in the very largest
specimens. Our single individual is larger
than any of the other specimens that we
refer to this species and we attribute the dif-
ferences in our specimen to its larger size.
Description of the soft parts. Incurrent
region with 7 papillae on either side, no
single tentacle; adductor muscles very small,
oval and dorsal in position, anterior adduc-
tor muscle somewhat larger than the poste-
rior; gills moderately large, 24 gill plates on
each side of tire axis; hind gut penetrates
slighth' into the mantle, ventral limit of
hind gut loop scarcely ventral to the ventral
edge of the anterior adductor muscle; vis-

typhlosole

posterior

adductor

muscle

digestive
gland

hind
gut

incurrent
papillae

anterior
sense
organ

palp

adductor

muscle

Figure 19. Tindaria cytherea (Dall). Semidiagrannmatic drawing of the body and mantle organs as seen
from the right side.

38 BuIJeti7i Museum of Comparative Zoology, Vol. 148, No. 2

rectal guides

anus /

excurrent
aperture

\ aperture

incurrent
aperture

Figure 20. Tindaria cytherea (Dall). Detail of the
posterior part of the left mantle margin.

ceral ganglion close to the posterior adduc-
tor muscle.

Remarks. Fortuitously, the soft parts of
our single specimen had become detached
from the shell on preservation. There are
no great differences in the morphology of
the soft parts from those of other species,
many being the consequence of the large
size of the specimen ( Fig. 19 ) . Hence, the
development and the extension of the inner
mantle fold to form a feeding aperture be-
low the incurrent region is particularly well
marked (Fig. 20). Similarly, the anterior
sense organ is well-developed. The gills are
large (Fig. 21) but not fused to form an
obvious pumping organ, although in life,

pallial Figure 21. Tindaria cytherea (Dall). Latero-ventral

muscles view of gill.

with the aid of peripheral interlocking cilia,
they could possibly function as such. The
gill axes which connect with the ventral
side of the excurrent region probably act as
guides during the expulsion of faeces from
the anus. The palps, probably because of
the large size of this specimen, have a
greater number of ridges ( 18) than in other
species of Tinclaria, but they are small in
comparison with other protobranch genera
of the same size. The palp proboscides are
extremely large and elongate. The mouth,
unlike the other species here described, is
close to the anterior adductor muscle. The
course of the gut is similar to that of T.
caUistiformis and T. hessleri. The stomach
is extremely large, with an extensive gastric
shield on the left dorsal side, and bears a
tooth close to the three apertures of the
digestive diverticula. There are 10 sorting
ridges on the right side (Fig. 22). The
ganglia are extremely large.

Tindaria miniscula, new species
Figures 23-28

Holotype: M.C.Z. 279901, from Atlantis 11 cruise

42, Station 197, in 4565 to 4595 m.
Depth range: 4559-4566 to 4612-4630 m.

Cruise

Station

No.

Depth

(m)

No. of
Specimens

titude

Longitude

Gear

Date

Atlantis

Atlantis
Atlantis

II
II

198

197

196

4559-4566

4565-4595
4612-4630

Angola B

27
2

isin

10
10
10'
10'
10'

=24.0'S-

=29.0'S

'29.0'S-

29.0'S

9'=04.0'E-

9°09.0'E

9°03.0'E-

9°04.0'E

ES
ES

21.5.68

21.5.68
21.5.68

Specific Description. Shell minute, swollen; beaks prominent and strongly
strong, elongate, oval in outline, with uni- curved medially; escutcheon forms a con-
form concentric ridges; umbo large and spicuous concavity in dorsal margin (Fig.

Deep Sea Protobranxhs • Sanders b- Allen

oesophagus

digestive
duels

sorting
area

Figure 22. Tindaria cytherea (Dail). Internal detail of the left and right sides of the stomach.

27c); external ligament elongate, narrow,
extending posteriorly to near the insertion
of the second tooth of posterior hinge plate
series and extending anteriorly to tlie inser-
tion of the first tooth of anterior series;
dorsal shell margin weakly convex; antero-
dorsal margin short, rather sti'aight and,
distally, sloping ventrally to form a smooth
curve with the anterior margin; postero-
dorsal margin longer, slightly convex to
form continuous curve with the posterior
margin; anterior and posterior margins
evenly rounded; ventral margin elongate
and moderately convex.

Hinge plate thin, rather weak with a rela-
tively broad edentulous space separating
the anterior and posterior rows of teeth;
posterior hinge series with six chevron-

shaped teeth; anterior hinge series shorter,
with three similar teeth ( Fig. 23) .

Tindaria miniscula differs from T. callisti-
formis, T. hessleri and T. cijtherea in its
small size, oval shape, few teeth on thin
hinge plate, the extensive endentulous
space on hinge plate beneath umbo and the
more medial position of the umbo.

Morpliology of the soft parts. Incurrent
region with three papillae to the left side
and two to the right; adductor muscles
small, unequal in size; gill very small with
six to seven plates to each demibranch; palp
with five to six ridges; hind gut penetrates
mantle of right side almost to the pallial
line; part of hind gut lies adjacent to the
posterior face of the anterior adductor

40 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Figure 23. Tindaria miniscula Sanders and Allen. Ex-
ternal view of the left valve and internal view of the
left and right valves.

muscle; visceral ganglion contiguous with
the posterior adductor muscle.

The two papillae to the left of the incur-
rent region alternate with the three to the
right so that the dorsalmost papilla is on the
right, i.e., the two most venti'al are on the

left. The third left ventral papilla may
possibly be homologous to the single nucu-
lanid tentacle but in form and histology it
cannot be separated from the others. A pair
of well marked faecal guides or ridges mark
the ventral limit of the excurrent aperture.
Below the incurrent region the extended
inner mantle folds overlap to form a feeding
aperture. Separation of the aperture is not
permanent. Inward of the feeding aperture
and the incurrent aperture is a band of very
large epithelial gland cells. Adductor mus-
cles are small and the posterior is larger.
The gills are very small, approximately
seven pairs of alternating gill plates set
tangentally across and behind the posterior
part of the body. The gill axes are attached
to the mantle edge at the level of the faecal
guides. There are a few muscle fibers in
the axes and two well-developed muscles
in the plate. The palp is relatively large,
extending approximately half the total
length of the animal; the palp proboscides
are stout ( Fig. 24 ) . The mouth is set poste-
rior to the anterior adductor muscle, the
oesophagus extends forwards, turning short
of the anterior adductor muscle, postero-
dorsally to the stomach. The lumen of the
oesophagus is exceptionally large, as is the
stomach. The latter occupies the bulk of
the body space. The stomach is almost
entirely lined with the gastric shield, and
is surrounded by a fine network of muscle
fibers. Diatom frustules are the main or-
ganic content of the stomach. It is possible
that the digestive gland is composed of two
regions, the umbo non-pigmented and the
remainder a pale brown color, but sections
indicate no obvious histological differences.
The hind gut forms a single loop to the
right-hand side of the body, penetrating
the lumen of the mantle and skirting the
anterior adductor muscle, but not reaching
the inner mantle lobe. At its maximum
point of penetration it lies some distance
dorsal to the anterior mantle sense organ.
The hind gut also extends deep into the
foot ventral to the pedal ganglia. A typhlo-
sole is present along the entire length of

Deep Sea Protobranchs • Sanders b- Mien 41

nd gut

oesophagus

anterior
sense organ

pedal ganglion

foot

Figure 24. Tindaria miniscula Sanders & Allen. Semidiagrammatic drawing of the body and mantle organs
as seen from the right side.

the gut. The foot is relatively small, and
placed far back on the visceral mass. It
bears a relatively large, triangular, poste-
riorly directed heel and a small 'byssal'
gland. The sole of the foot is small, divided
and peripherally papillate with glands to
the inside of the papillae. The ganglia are
large, particularly the pedal, the visceral
and cerebral ganglia are attenuate pear-
shape. There is a pair of small but typical
protobranch statocysts dorsal to the pedal
ganglia. The kidney is veiy small; sexes
are separate, whole mounts showing initial

stages in ovarian development with approx-
imately 12 ova lateral to the stomach.

Although the total number of specimens
in our samples is small, and the size range
is so much smaller than in other species
(0.8 to 2.5 mm), the population histogram
remains clearly skewed to the left as it is in
Tindaria caUisfifonnis (Fig. 25). Similai-ly,
the height/length ratio shows a wide range
of values varying from 0.66 to 0.78 over the
length range (Fig. 26). However, we may
not have sampled the smaller juvenile
stages which might have passed tluough the

10-1

STA. 197 a 198

LENGTH (mm)

Figure 25. Tindaria miniscula Sanders & Allen. Size frequency histogram of specimens collected at Sta-
tions 197 and 198.

42 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

90-

80-

° □

D
D

70-

D D dQ on

60-

•

50-

• •

•
•

•

•
•

•

•
•

•

40-

1.0

1.5
LENGTH (mm)

2.0

Figure 26. Tindaria miniscula Sanders & Allen. Graph showing height/total length (O) and umbo to poste-
rior margin total length {•) plotted against total length.

0.42 mm apertures of our screen. The anterior as the size of tlie shell increases

innbo, unlike other species of Tindaria, is (52 per cent to 57 per cent), i.e., there is an

almost central in position in the smallest increase in length of the posterior part of

specimens and becomes progressively more the body with increasing age (Fig. 28).

A ^' B ^ C

Figure 27. Dorsal views of A, Tindaria callistiformis; B, Tindaria hessleri; C, Tindaria miniscula.

Deep Sea Protobranchs • Sanders h- Allen 43

Family INCERTAE SEDIS^
Pseudotindaria new genus

Type species: P. erebus (Clarke, 1959)

Shell stout, oval in outline, inequilateral
and with strong, uniform, concentric ridges;
external Hgament extends anterior and

1 Considering the present systematic state of the
protobranchiate bivalves and the need to erect a
number of additional families to include the diver-
sit>- of morphologies, we feel that it is premature
at this time to assign the genus Pseudotindaria to
a presently defined family.

posterior to the umbo; hinge plate strong
and continuous below umbo, with at most
a minute endentulous area between the
anterior and posterior hinge teeth series;
siphons present and fused ventrally; si-
phonal embayment shallow; gills horizontal
to the anteroposterior axis, or nearly so;
palps moderately large with many narrow
ridges on the inner faces; mouth opens close
to the anterior adductor muscle; hind gut
coils on both sides of the body and does not
penetrate the mantle.

Tindaria

Pseudotindaria

1 ) Roimded, robust shell concentrically hned 1 )

2) Hinge line strong, supporting a series of well- 2)
dexeloped teeth, continuous below the umbo

3) Ligament external 3)

4) Tentacle present or absent 4)

5 ) No siphons, fringing papillae around inhalent 5 )
region

6 ) Palp with iew ridges 6 )

7 ) Relatively few gill plates 7 )

8) Hind gut with tviihlosole, single loop to the 8)
right of body penetrating the mantle

9) 'Byssal' gland small and ovoid 9)

10) Ganglia very large, pedal ganglia in a median 10)
position within foot

Same
Same

Same

Tentacle present

Siphonate, without papillae

Palp with many ridges
Relati\ely many gill plates
Hind gut without typhlosole, loops and/or
coils to right and left of body crossing be-
fore and behind the stomach, does not
penetrate mantle

'Byssal' gland very large and cylindrical
Ganglia small, pedal ganglia in upper half
of foot

Pseudotindaria erebus (Clarke, 1959)
Figures 29-40

Tindaria crchus Clarke, 1959. Proc. Malacol. Soc.
London, 33: 236. Text. fig. 1(1). (Type locality:
m V Thcia, Station 9, Lat. 31°42'N, Long. 68°
OS'W; type specimen, Mus. Comp. Zool., No.
218182.)

Neilonella galathea Knudsen, 1970. Galathea Rep.,

XI, p. 58. Text fig. 38, B-D; Plate 5, fig. 2,

Plate 6, fig. 6. (Type localit>-: Galathea Station

66, Lat. 4°00'S, Long. 8°25'E; t\pe specimen,

Zool. Mus. Univ. Copenliagen. )
Pre\ious records: Single station west of Bennuda

at 31°42'N, 68°08'W in 5203 m (Clarke, 1959).

Single station off W. Africa 4°00'S, 8°25'E at

4018 m depth (Knudsen, 1970).
Present record: Deptli range = 2644-2754 to

5007 m.

Cruise No.

Station
No.

Depth

(m)

No. of
Specimens

Latitude

Longitude

Gear

Date

North America Basin

Chain 50
Atlantis II
Atlantis II

24
17

123

3834
4853
5007

1 37°59.2'N

2 34°39.0'N

Canaries Basin

69°26.2'W
64°14.0'W
66°56.0'W

ES
ES
ES

5.7.65
22.8.66
14.12.65

Discovery

6714

3301

1 27°13'N

i5°4rw

20.3.68

44 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Cniise No.

Station
No.

Depth

(ni)

No. of
Specimens

Latitude

Longitude

Gear

Date

Cape Verde

Basin

Atlantis II
Atlantis II

31
31

148

149

3814 3828
3861

2
12

10°37.0'N
10°30.0'N

18n4.0'W
18°18.0'W

ES
ES

7.2.67
7.2.67

Brazil Basin

Atlantis II
Atlantis II

31
31

156
155

3459
3730-3783

Guiana B

00°46.0'S-

00°46.5'S

00°03.0'S

isin

29°28.0'W-

29°24.0'W

27°48.0'W

ES
ES

14.2.67
13.2.67

Knorr 25
Knorr 25
Knorr 25

307
288

287

3835-3862
4417-4429
4934-4980

Angola B

12°35.4'N-

12°40.8'N

1P02.2'N-

11°03.8'N

13°16.0'N-

13°15.8'N

isin

58°59.3'W-

59°09.2'W

55°05.5'W-

55°04.8'W

54°52.2'W-

54°53.1'W

ES
ES
ES

3.3.72
25.2.72
24.2.72

Atlantis II
Atlantis II

42
42

200
195

2466-2754
3797

36
145

9°41.0'S-
9°43.5'S
14M0.0'S

10°55.0'E-
10°57.0'E
9°54.0'E

ES
ES

22.5.68
19.5.68

Description of Shell. Tlie specimens in
our collection depart from Knudsen's ( 1970)
description only by having a very narrow
and not a 'rather wide' edentulous space
separating the anterior and posterior hinge
teeth series. Tliey differ from the specimen
described by Clarke ( 1959 ) by having one
more tooth in both the anterior and poste-
rior hinge series.

With the additional material from our
collections we can add further obsei-vations
to the precise descriptions given by Clarke
(1959) and Knudsen (1970) (Figs. 29,
30 & 31). Shell stout, with strong, uni-
form concentric ridges, oval in outline and
slightly extended posteriorly; umbones low,
anterior in position; beaks not prominant,
curved medially and slightly anteriorly; es-
cutcheon forms a shallow concavity; exter-
nal ligament elongate, narrow, extending
posteriorly in escutcheon to about insertion
of sixth or seventh tooth on the posterior
hinge plate and anterior to the third tooth
of the anterior hinge plate; hinge plate
strong and moderately thick with a minute
edentulous gap immediately below the um-
bonal beak; posterior hinge plate with about
14 teeth, medial teeth small, and on the dor-

sal side of the hinge plate, distally the teeth
enlarge the bases nearly spanning the width
of the hinge plate; anterior hinge plate with
about 11 teeth, medial three or four teeth
minute, rod-shaped and restricted to the
upper half of the hinge plate; distally the
teeth enlarge becoming chevron-shaped and
rather stout; postero-dorsal shell margin
slightly convex forming a smooth curve with
the posterior margin; antero-dorsal margin
short but longer than in T. callistiformis or
T. hessleri, margin straight medially or even
slightly concave, anteriorly convex fomiing
a continuous curve with anterior end; ven-
tral margin long and relatively convex.

Morphology of the soft ports. Siphons
are developed posteriorly from the inner
muscular fold (Figs 32 & 33). They are
fused dorsally but not ventrally so that the
lumen of the excurrent siphon is not sepa-
rated by tissue from the incurrent, nor is
the latter separated by tissue fusion from
the mantle gape below. Central and venti^al
ridges are present and when these are ap-
posed they effectively separate the lumen
of the excurrent and incurrent siphons as
well as the feeding aperture below. A
single tentacle is inserted below the incur-

Deep Sea Protobranchs • Sanders 6- Allen 45

Figure 29. Pseudotindaria erebus (Clarke). Internal
view of left valve of type specimen No. 218182 Mu-
seum of Comparative Zoology, Harvard.

Figure 28. T/ndana m/n/'scu/a Sanders & Allen. Growth
series in lateral view.

rent aperture on the right side and, histo-
logically, it is similar to that of Tindaria.
A food aperture is also developed from the
hypertrophied inner muscular fold giving
rise to a third, but smaller, channel. The
anterior sense organ is well-developed. Ad-
ductor muscles are oval, although not
greatly elongate, and situated at a relatively
' greater distance in from the shell margin
than is the case in Tindaria. Although tliere
is no marked development of the siphonal
, embayment, the siphons can be retracted
within the valves; the retractor muscles are

Figure 30. Pseudotindaria erebus (Clarke). Internal
view of left and right valves of specimen from Station
195.

46 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

Figure 31. Pseudotindaria erebus (Clarke),
outline; C, dorsal view of entire shell.

A, anterior view of shell in outline; B, dorsal view of valve In

not particularly well-developed. There is
no great concentration of gland cells inter-
nal to the feeding aperture; although there
are small mucous cells lining the main man-
tle rejectory ti'act leading to this area.

The gills are more or less horizontal with
14 to 16 plates on each side of the axis, the
plates on either side alternating. The gill
plates are approximately equal in size, those
of the inner demibranch slightly larger than
those of the outer, particularly posterior
to the body; each has a fan of three
muscles in die transverse plane which pene-
trate the axis as a retractor muscle, and on
either side of the retractor muscle are
longitudinal muscles running the length of
the axis (Fig. 34). There is no tissue fusion
to the mantle and/or the body opposite to
form a diaphragm because the separation
of the hypobranchial cavity from the re-
mainder of the mantle cavity is accom-
plished by ciliary junctions. Note that the

cerebro-visceral connectives lie close to the
junction of the gill axis with the body.

The palps are relatively larger than those
of Tindaria with many more ridges (17 to
30 ) . The mouth is close to the anterior ad-
ductor muscle and the palps extend more
than half way across the body. The palp
proboscides are relatively large. The foot is
moderately large widi a well-defined neck;
the fringing papillae and heel are very
small. A large 'byssal' gland is present, the
hyaline central portion of which is elongate,
cylindrical and different in shape from any
of those described to date (Sanders and
Allen, 1973; Allen and Sanders, 1973) (Fig.
35). As in other genera, there is a centi'al
sagittal strip of tissue dividing tlie gland in
half, the gland opening just posterior to the
sole of the foot.

The oesophagus extends dorsally, and
posteriorly, opening into a relatively simple
stomach which has a few low-crested sort-

Figure 32. Pseudotindaria erebus (Clarke),
seen from the right and left sides.

Semidiagrammatic drawings of the body and mantle organs as

Deep Sea Protobranchs • Sanders ir Allen 47

posterior
adductor
muscle

kidney —

48 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

hindgut

excurrent
siphon

incurrent
siphon

tentacle

siphona
retractor
muscles

Figure 33. Pseudotindaria erebus (Clarke). Detail ot
siphonal region.

hind gut

hypobranchial

gland "~~>*£iiA

gill
axis

muscle

mantle

Figure 34. Pseudotindaria erebus (Clarke). Trans-
verse section through gill and mantle to show mus-
culature and glands.

Figure 35. Pseudotindaria erebus (Clarke). Trans-
verse section through the foot to shovj the position of
muscles and gland.

ing ridges on tlie right side (Fig. 36). The
long axis of the stomach and style sac is
oblique to the body axis and directed pos-
tero-ventrally. The hind gut does not pene-
trate far into the foot before turning dorsally
to the left ventral side of the body. Tliere-
after it passes to the right side of tlie body
in front of the stomach (posterior to the
oesophagus) forming a small loop before
returning to the left where it outlines the
periphery of the stomach and the digestive
gland (Fig. 37). After forming this single

hrndgul

gastric
sh.eld

aperfufe to

digestive duel

Figure 36. Pseudotindaria erebus (Clarke). Trans-
verse section through body to show stomach and
outline of hind gut sections.

Deep Sea Protobranchs • Sanders ir Allen 49

Figure 37. Pseudotindaria erebus (Clarke). Dorsal
diagrammatic view of stomach and hind gut.

coil it returns to the right side of the body
passing dorsal to the style sac and the ante-
rior part of the hind gut. Here it forms a
second loop to die outside of the first before
continuing to the anus via the mid-dorsal
line. The second loop on the right side and
the single coil on the left are appro.ximately
the same diameter and pass close to the

Figure 39. Pseudotindaria erebus (Clarke). Growth
series in lateral view.

posterior face of the anterior adductor mus-
cle. No typhlosole is present nor does the
hind gut penetrate the mantle.

The ganglia are not particularly large, the

2.0

Immature

1 STA. 200

35 Specimens

10-

_ ^^ ■ _

1 STA. 195

145 Specimens

10-

1 ^ ■_ _■ ■ ,

■ _J ■

^^^^^^^^^ ^\

3.0

4.0

LENGTH (mm)

Figure 38. Pseudotindaria erebus (Clarke). Size frequency histograms of samples from two stations. The
dashed lines indicate the size at which gametogenesis is evident.

50 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

80-

60-

n
o D

.♦

D D

D " n

• • •

1 T

3
LENGTH (mm)

Figure 40. Pseudotindaria erebus (Clarke). Graph showing height/total length (□) and umbo to posterior
margin total length (•) plotted against total length.

visceral and cerebral are elongate, the pedal
ganglia are positioned high in the foot, the
visceral ganglia abont the posterior adduc-
tor muscle ( Fig. 32) . Note that the foot can

Figure 41. Pseudotindaria championi (Clarke). Inter-
nal views of left valve of type specimen No. 224957
Museum of Comparative Zoology, Harvard with en-
larged detail of the umbonal region.

be extended backwards far into the poste-
rior part of the mantle cavity and, as in
other protobranch species, may well assist '
in pushing rejected faecal material out of ■ j
the mantle cavity.

Size and Reproduction. The population
histograms of stations 195 and 200 ( Fig. 38)
show that the skew to the left is not nearly
so marked as it is in T. callistiformis. Spec-
imens less than 2.5 mm in total length are
all immature and those 3 mm or more show
signs of gonad development. Sexes are
separate, the sex ratio is even. Maturity
increases with increasing size. A fully ma-
ture female measuring 6.0 mm total length
contained approximately 800 ova having a
maximum length of 142 ^u,. The gonads
initially develop at the ventral perimeter of
the body to the inside of the outer hind gut
loop. Eggs mature simultaneously. On an
average, 42 per cent of a given population is
maturing, samples ranging from 33 per cent
to 49 per cent.

As in T. callistiformis, there is a relatively
high degree of variation in shell dimensions,
particularly in respect to the height/total
length ratio, which varies from 0.71 to 0.87.
There is no relation between this variation
and the increasing size of the shell. The lat-

Deep Sea Protobranchs • Sanders b- Allen

ter also applies to the total length umbo-
posterior margin length ratio. The umbo,
although anterior, is not so far forward as
it is in T. callistiformis (51-63 per cent
of the total length). There is a slight
trend for the posterior umbonal length to
increase with increasing size although the
\'ariation remains wide throughout the size
range (Figs. 39 and 40). Maximum total
lensfth recorded is 6.0 mm.

Pseudotindaria championi (Clarke,
1961)
Figures 41-43

Timlaria championi, Clarke, 1961. Bull. M.C.Z.
125, 372. Plate 2, figs. 1 and 4. (Type locality:
r/v Vema, Biology Station 12, Lat. 38°58.5'S,
Long. 41°45'W, type specimen M.C.Z. 224957).

Previous records: Depth range = 3116 to 5133 m.

Argentine Basin — 1 station. Ref. Clarke, 1961.

Cape Basin — 1 station. Ref. Clarke, 1961.

Present records:

Cruise

Station
No.

Depth No. of
( m ) Specimens

Latitude

Longitude

Gear

Date

Argentine Basin

Atlantis 11

269A

3305-3317 5

37°13.3'S

52M5.0'W

26.3.71

Atlantis 11

242

4382-4405 3

38°16.9'S

51°56.rW

13.3.71

Description of SJwU. Since we are able
to add a number of further observations to
Clarke's ( 1961 ) very adequate description
of the hard parts of Pseud of i nil ario cham-
pioni, a fairly extensive redescription fol-
lows.

Shell stout, with strong concentric ridges,
subovate in outline, and slightly extended
posteriorly; umbos low, anterior in position,
beaks moderately prominent and curv^ed
medially; escutcheon forms a shallow con-
cavity; external ligament elongate, narrow,
extends from about the insertion of fourth
tooth on the posterior hinge plate to about
second tooth of the anterior hinge plate;
hinge plate strong and somewhat angular
with moderately extensive edentulous gap
below umbo; posterior hinge plate with
about eight chevron-shaped teeth medial
three teeth becoming progressively smaller
and restricted to upper margin, distally
teeth enlarge in size wnth bases almost
spanning the width of the hinge plate, dis-
talmost teeth rather massive; anterior hinge
plate with about seven teeth, medial tooth
small, rvidimentary and confined to middle
of hinge plate, more distal teeth chevron-
shaped and progressively larger inserted
progressively further from the dorsal shell
margin; entire shell margin smoothly
rounded in outline; postero-dorsal shell
margin much more convex than short, al-

most straight antero-dorsal margin; poste-
rior margin broad; ventral margin long and
moderately convex. Total length of 3.56
mm. Pseuclotindaria championi is readily
distinguished from P. erehus by its more
rounded outline, a more medial positioning
of the umbo and its fewer teeth.

Morphology of the soft parts. The mor-
phology of the animal is very similar to that
of P. erehus; combined incurrent and excur-
rent siphons are present, the respective
channels being separated by approximation
of the tissues along the length of the mid-
line of the siphon. Similarly the incurrent
siphon is separated from the pedal gape by
apposition of the right and left ventral
edges. There is a shallow embay ment
formed by the extension of the mantle edge
between the insertion of the pallial mus-
cles and the outer part of the sensory lobe.
Inserted on the right-hand side at the ven-
tral limit of the embayment is a small
single sensory tentacle (Figs. 42, 43). The
food apertiu-e is not very clearly defined
in P. championi, and barely separates from
the extensi\'e pedal gape — its ventral limit
is marked by the posterior edge of a well-
marked ridge of glandular tissue to the in-
side of the inner muscular lobe of the mantle
edge in the posterior half of the pedal gape
(Fig. 42). A pair of well-developed ante-
rior mantle sense organs are present below

52 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

digestive
diverticula

posterior

adductor

muscle

timd
gut

oesophagus

Deep Sea Protobranchs • Sanders 6- Allen EC

Figure 43. Pseudotindaria championi (Clarke). Lat-
eral view of the course of the hind gut as seen from
the right side. Stippled sections are positioned on the
left side of body, blacked sections on the right.

f the anterior adductor muscle. The adduc-
tor muscles are oval, equal in size, with the
'quick' and 'catch' portions clearly seen.

The 2;ills are horizontal, each with 10 or
11 pairs of plates. The gill axis extends
be)'ond the posterior plates to fuse with the
siphonal tissue at the junction of incurrent
and excurrent channels, and as in T. erebus
there is no fusion between gill and mantle,
body or opposite gill. It seems likely that
I as in other protobranchs, tlie posterior limits
■ of tlie gill axes act as guide rails to facili-
tate the removal of bulky faecal rods. The
palps are moderately large and extend from
the mouth partly across the foot; the dorsal
limit of the ridged area is attached to the
body in front of the junction of the anterior
edge of the muscular part of the foot to the
viscera. The many ridges (approximately
25) spread fanwise, posteriorly. The re-
tracted palp proboscides do not extend be-
yond the posterior margin of the foot. The
foot is dorso-venti'ally elongate, the sole is
not exceptionally large, the tip is pointed,
with the fringing papillae moderately small,
rounded and low crowned. The heel is pro-
duced as a small, short process with a mod-
erately large 'byssal' gland, similar to that
described for P. erebus. The gut is also
similar to that of P. erebus (Fig. 42),
with tlie hindgut arranged in the same con-
figuration. However, the relative diameter

Figure 44. 'Tindaria' acinula Dall. Internal views of
the left and right valves.

of the hind gut is greater in P. championi,
in which the gut occupies much more of the
body space (Figs. 32, 42).

DISTRIBUTION PATTERNS

Of the six species considered in this paper,
four members of the genus Tindaria and
two representatives of the genus Pseudo-
tindaria, two are cosmopolitan while the
remaining four appear to be confined to
restricted regions of the Atlantic. The two
widely distributed species Tindaria cal-
lisfiforniis and Pseudotindaria erebus are
abyssal species and have been collected
from the Nortli America, Guina and Angola

Figure 42. Pseudotindaria championi (Clarke). Semidiagrammatic drawings of the body and mantle organs
as seen from the left and right sides.

54 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

hind gut

anterior
adductor
muscle

tenlocle

Figure 45. 'Tindaria' acinula Dall. Semidiagrammatic
drawings of the body and mantle organs as seen from
left and right sides.

Basins. In addition, P. erehus has been
taken in the Canaries, Cape Verde and
Brazil Basins and T. caUistiformis in the
Argentine Basin. Pseudotindaria erehus,
which occurs in shallower depths, has been
found in about 2650 m in the Angola Basin,
but T. callisfiformis has never been col-
lected shallower than about 3300 m. The
only region so far examined where P. erehus
has not been obtained is the West Europe
Basin. However, the samples analysed to
date from that region have been limited to
depths less than 2380 m. Similarly, on the
basis of depth distribution of T. calUstifor-
)nis, there are only four stations of appropri-
ate depths where it has not been found
(two stations in the Angola Basin and two
stations in the Brazil Basin). Probably
these forms are present throughout the At-
lantic wherever there are adecjuate depths.
Oiir data suggest that Tindaria caUistiformis
may be numerically significant only in a
small subset of its vertical and perhaps its
horizontal range. In most samples where it

is present, T. caUistiformis makes up but a"(
minor fraction of the protobranch fauna,
0.08 to 1.97 per cent at ten stations 4600 m
or deeper. At the shallowest station, in 3305
to 3317 m of water, it constitutes 4.48 per
cent. In the remaining six samples, taken in "
3806, 3828, 3834, 3906 to 3917, 4417 to 4429,
and 3825 to 3862 m, this species formed
20.84, 18.99, 13.80, 27.61, 3.82 and 0.74 per
cent respectively of the protobranch spec-
imens. The first three samples were taken
from the North American Basin, the fourth
was from the Argentine Basin, and the re-
maining two, from the Guiana Basin.

Despite its broad horizontal distribution,
Pseudotindaria erehus must be a very rare
animal at least in part of its range. Of more
than 22,500 protobranchs collected from the
Gay Head-Bermuda transect in the North
America Basin, only four specimens of P.
erehus have been taken. Alternatively, in
the Angola Basin it may be numerically im-
portant, 19.05 per cent at station 200 and
20.03 per cent at station 195. The values
found in the Canaries, Cape Verde, Guiana,
Brazil and Argentine Basins fall between
these extremes.

Of the remaining species, Tindaria hes-
sleri appears to be a lower slope-upper
abyssal rise inhabitant of the northeast At-
lantic. We have taken it at a single station
in the West Europe Basin and from five of
the six stations in its depth range in the
Cape Verde Basin. Thus it appears to be a
constant faunal constituent within the ap-
propriate depth range and in the latter
basin comprises, numerically, nearly one to
ten per cent of the protobranch bivalves.

Tindaria cytherea has a narrow depth
range, restricted to intermediate slope
depths in the northwest Atlantic. It has so
far been reported from a few localities in
the North America, Gulf of Mexico and
Caribbean Basins. We have collected but a
single specimen in 1000 m depth off Ber-
muda. It is probably not present at equiv-
lent depths at the New England end of the
Gay Head-Bermuda traverse, which have
been extensively sampled.

Deep Sea Protobranchs • Sanders «L~ Allen 55

Tseudotindaria championi is an abyssal
ipecies of the high southern latitudes known
rem three localities in the Argentine Basin
ind a single sample in the Cape Basin.

The remaining species, Tindaria minis-
cula, is probabh' limited to the deeper
abyssal depths of the southeast Atlantic.
We have found it at three stations in the
Angola Basin where it formed 0.99, 2.04
and 3.29 per cent of the protobranch fauna.

iPALEONTOLOGICAL RELATIONSHIPS

Pojeta (1971) has pointed out that all
known 'nuculoids" from the early Paleozoic
(Ordovician) lacked a resilifer but had in-
stead a hinge plate continuous beneath the
umbo. In addition, these forms possessed
a continuous row of teeth rather than teeth
that were separated into an anterior and
posterior hinge series. Among the living
protobranchs, the genera Tindaria, Tseudo-
tindaria and Neilonella have a similar com-
bination of morphological characters.

Most Ordovician nuculoids ha^'e been
placed in the extinct family Ctenodontidae.
However, the careful studies of Pojeta
( 1971 ) clearly show that the Ordovician
nuculoids were a "highh' ^'aried and suc-
cessful group," a fact masked by the pro-
nounced conservatism of the systematics.
Earlier, \^errill & Bush (1897) noted that
' the ctenodont PaJaeoiieiJo "agree in nearlv
, all essential characteristics with the living
genus Tindaria." \\'ithin the complex of
Ordovician ctenodont shell morphologies
cited by Pojeta ( 1971 ) can be found appar-
ent homologues of Tindaria and Tseudotin-
daria; e.g., Talaeoneilo fecunda (Hall), and
XciloneUa; e.g., Decaptrix aff. D. harts-
villensis (Stafford) and Decaptrix baffin-
ense (Ulrich).

In the present paper we have documented
, the profound differences in the soft part
' anatomy of Tindaria and Tseudotindaria,
two genera which have almost identical
shells. Neilonella is more elongate and ros-
trate but the soft part anatomy resembles
that of Tseudotindaria, e.g., they have si-

phons, many gill plates and palp ridges and
lack posterior papillae. Thus, these fev/
modern genera also represent a varied com-
plex of morphologies. Yet, on the basis of
shell morphology, there is no reason why
these living genera sh(nild not be the mod-
ern descendants of the Paleozoic ctenodonts,
the oldest lineage of protobranchs with
typical chevron-shaped teeth, representing
a group or groups of bi\'alves with remark-
ably conservative shell features.

In comparison, we have pointed out else-
where that the modern nontaxodont proto-
branchs such as the families Nucinellidae
(Allen & Sanders, 1969), Lametihdae and
Siliculidae (Allen & Sanders, 1973) are
analogs of the Palaeozoic actinodonts and
may, indeed, be direct derivatives.

DISCUSSION

The present paper, in addition to defin-
ing a moiphology that stands apart from all
other protobranchs, emphasizes more clearly
than in any other group of the Proto-
branchia the danger of relying on shell
character to define a taxon. Thus, Tindaria
and Tseudotindaria on shell characters alone
would have been (indeed were) placed in
the same genus, but the soft parts show
that they are greatly different. This has the
unhappy consequence that anatomical stud-
ies must be made to confirm the placement
of many of the species referred to the genus
Tindaria.

Tseudotindaria is difficult to place in
relation to other known protobranch genera
at this stage in our researches. Some species
of Neilonella described by Knudsen (1970)
certainlv resemble Tseudotindaria and it is
our intention to analyze the "Neilonella
complex" in a subsequent paper of this
series. Similarly, so-called Tindaria acinula
( a species that occurs in our samples ) is an
example of a species wrongly described in
the literature as a tindariid which, like
Tseudotindaria, has siphons, not tentacles,
and a large palp with many ridges. The
hind gut does not penetrate the mantle and

56 Bulletin Mtisciim of Comparative Zoology, Vol 148, No. 2

the large posterior adductor is not close to
the shell margin (Figs. 44, 45). However,
unlike Fseudotindaria, the hind gut crosses
to the left side of the body in front of the
mouth and there forms four coils, and the
valves are posteriorly rostrate rather than
oval. Again, this species will be considered
in a later paper.

The species of Tindaria are characterized
b\' tlie lack of posterior mantle fusion and,
in this sense, as well as in liaving a gill
placed obliquely across the mantle cavity,
they are at first sight akin to the Nuculoidea.
We hypothesize that the tindariids may
have similar habits to some species of Nu-
cula (see below). Nevertheless, the poste-
rior mantle edge is highly specialized and,
in addition to the pedal gape, it fomis
functional posterior incurrent, excurrent
and feeding apertures. The apertures are
formed by the apposition or overlapping of
the inner and, sometimes, middle lobes of
the mantle. The excurrent regicm is partic-
ularly circumscribed, the space between the
adductor muscle and the first papillae being
only wide enough to allow the passage of
the faecal rod. Knudsen (1970) reports a
simple flap-like extension of the mantle on
the dorsal side of the aperture in two Pa-
cific species. In T. bengaJensis and T. com-
pressa, at the ventral limit of the excurrent
region and extending across the mantle
edge on each side is a shallow ridge which
might possibly be considered to be the first
stage in the development of a siphon, to
which the attenuate part of the gill axis
attaches. The anus opens opposite this
channel. In many specimens the dorsal-
most papilla on each side of the incurrent
region point inwards, parallel to tliis chan-
nel. The number of papillae on either side
of the aperture varies between species. An
unpaired papilla or tentacle, which in some
cases is longer than the other papillae, may
be present on the right or left side at the
ventral limit of the inhalent region. This
may or may not be equivalent to the un-
paired siphonal tentacle of other nuculanids.
Histologically there is no difference between

the two. The papillae consist of a centi-ali^
pair of longitudinal muscle bands with a i '
haemocoele between the muscles and the
outer epithelium. There are no major
nerves supplying the papillae and inner\'a-
tion is probal)ly in the form of a number of
fine fibers. There are also no concentrations
of gland cells, thus the papillae are in no ;
way comparable to the feeding tentacles of
the deep sea carnivorous Verticordiidae
(Allen & Turner, 1974). We can only as-
sume that the papillae have a generalized
sensory function and possibly act as a
coarse filter for the incurrent flow gene-
rated by the pumping action of the gills.

We have been able to keep T. callistifor-
mis alive for approximately one month. The
soft parts are virtually colorless in life. Al-
though the metabolism of these animals was
clearly affected by their long journey to the
surface, to reduced pressure and through
the varying changes of temperature of the
water column, their condition was suffi-
ciently good to confirm tliat the papillate
region defines the incurrent area and that
the palp proboscides are extended from the
feeding aperture. The inner mantle folds
are extended beyond the shell margin in this
region. Although gross movements of the
gills occurred, no definite pumping rhythm
was observed. Ciliary movement was par-
ticularly languid and we hesitate to say
more than that the ciliary patterns within
the mantle cavity are similar to those de-
scribed by Yonge (1939) for more shallow-
living nuculanids.

The stomach takes up much of the body
space and it may be that the extension of
the hind-gut into the mantle is in part re-
lated to the lack of space between the body
wall and the stomach. Undoubtedly the
refractile nature of the food (diatom frus-
tules, etc.) to enzyme action is correlated
with the considerable extension of the gut
in deep-sea deposit feeding bivalves (Allen
& Sanders, 1966; Allen, 1971, 1973). One of
the many strategies used by the deep-sea
protobranch for housing the hind gut is
exemplified in the tindariids by their utiliza-

Deep Sea Protobranchs • Sanders h- Allen 57

Ition of mantle space. Exploitation of mantle
space occurs in other bivalves, as for the
gonads of mytilids — but deep-sea proto-
branchs are unique in the accomodation of
the hind-gut in this manner. There is much
e\'idence that material entering the stomach
in deep-sea protobranchs is less rigorously
sorted than that in shallow-water species.
Thus the ciliated sorting ridges are reduced
in number and size. The large tooth of the
gastric shield deserves mention because it
presumably does not act in the same man-
ner as in the Lamellibranchia. The material
coming forward from the style sac in the
protobranchs is a soft particulate mixture
and not a hard crystalline style. Hence it is
difficult to imagine that the tooth either
serves as a stop to the forward movement of
the style or as a cutting surface which
'turns' the head of the style with subsequent
release of style material. We believe that it
mav act here as a barrier to the encroach-
ment of space immediately posterior to the
oesophageal aperture by the style and/or
acts as a 'breakwater' in front of the ducts
to the digestive diverticula preventing ma-
terial from being forced into the ducts. It
may also act as a baffle to enhance the mix-
ing of the stomach contents in a fashion
analogous to the blade in a cement mixer.
There is evidence that selection of particles
does occur because a high proportion of the
stomach contents consists of diatom frus-
tules.

We believe that it is possible to deduce
something about the habits of the genus
Tinchiria. The strong robust concentrically
ornamented shell and rounded shape, to-
gether with an attached growth of hydroids
on many of the specimens, indicate a surface
dwelling rather than the infaunal habit for
the genus. The presence of tentacles around
the inhalent region and a pumping gill can
also be correlated to a stationary habit
close to the surface. The foot is by no
means as highly developed as in the smooth,
glossy, thin-shelled genera without epifauna
such as Siliciila and Spinula, which have all
the hallmarks of fast-burrowing infaunal

bivalves (Allen & Sanders, 1973). Never-
theless, the foot is moderately developed
and, in addition to any cleansing function,
it may well be involved in occasional rapid
movement, necessitated by predators or for
reorientation.

Tindaria species do not show an inverse
relationship between the size of the palp
and the size of the gill with increasing
depth, first shown in Abra (Allen & San-
ders, 1966) and later for some genera of
Protobranchia (Allen, 1973). We think
that a possible explanation is that gill size is
critical in relation to efficient pumping ac-
tion. Elsewhere (Allen & Sanders, 1966)
we have also shown that hind gut length is
much greater in deep water species as com-
pared with related shallow water forms. In
this, Tindaria is more consistent with the
general pattern. Thus, estimations of hind
gut volume per unit animal volume shows
that gut volume is half as large again in T.
caUistiformis ( 3305 to 5042 m ) as compared
with T. hessJeri (1739 to 2339 m). How-
ever, r. minisculu (3797 to 4566 m) is an
exception for it has a similar gut volume/
animal volume ratio, as in the two shallower-
dwelling tindarid species (see below). This
anomaly may be explained by the small size
of T. miniscula as compared with the other
species. Perhaps of more significance is the
fact that the gut volume of Pseudotindaria
(2644 to 5007 m) is similar to that of Tin-
daria, i.e., that irrespective of the great dif-
ference in hind gut configuration the volume
is not significantly different in the two gen-
era.

Gut volume/unit shell volume

cijdierca

1000 m

1.005

hessJcri

1739-2339

0.903

miniscula

3797-4546

0.954

callistifoiDiis

3305-5042

1.413

clianipiuni

3305-4405

erehus

2644-5007

0.915

The reproductive strategies of the genera
Tifidaria and Pseudotindaria are greatlv dif-
ferent. Pseudotindaria follows the typical

58 Bulletin Museum of Comparative Zoology, Vol. 148, No. 2

deep-sea pattern of approximately 50 per
cent of the population maturing at any one
time. The largest animals are the most
mature and the sex ratio is 1:1 (Sanders &
Allen, 1973; Allen and Sanders, 1973; Schel-
tema, 1972). (It must be noted that both
dioecious and hermaphroditic species occur
in the deep sea.) Our collections of Tin-
daria caUistiformis differ in the extremely
low percentage of the population that was
matming and also in the male-female ratio
of 5:1. However, it appears that the largest
animals are the most mature. We have
argued elsewhere (Sanders & Allen, 1973)
that, in the sparsely populated abyss, it is
an advantage to have large numbers of
sperm produced (and have a free plank-
tonic larva, even though of a short dura-
tion). We may be observing, in this case,
an exceptional condition even for Tindaria
and if so we have fortuitously encountered
a remarkably successful and isolated larval
settlement. However, there are reasons to
doubt this; not only is the pattern main-
tained at three widely spaced stations but
also the analysis of the T. miniscida spec-
imens indicates that this species is showing
a similar population structure, i.e., that these
are features common to the genus. It is
our intention to re-investigate these stations
at a different time of year following an
interval of some years to see whether the
structure has been maintained. More strik-
ing and less easily explained is the small
number of eggs present in the o\'ary, which
conflicts with the dominance of this species
in the samples and the large number of
small specimens in the sample.

SUMMARY

A new family of the Protobranchia, the
Tindariidae, is erected and included in the
Order Nuculanoidea. As of present, the
family is monogeneric; previously associated
genera, namely Pseudoglomus, NeUoneUa
and Mcdietia are not included because of
their markedly different morphologies.

Two new species are described — Tindaria
hessleri and T. rniiii.scula.

The species of Tindaria described here
show a number of common anatomical fea-
tures that are of particular note.

1) The posterior mantle edge bears
papillae on either side of the non-siphonate
incurrent aperture, the number varying ac-
cording to the species and to the size of the
animal.

2) The single loop of the hind gut and
associated visceral tissue penetrates the
right mantle to a greater or lesser extent. In
T. caUistiformis the loop approaches close
to the mantle margin immediately above the
anterior mantle sense organ while in T.
cytherea it barely penetrates the mantle.

3) Palp ridges are few in number.

4) Gametogenesis was observed in less
than 5 per cent of the specimens of T. cal-
listiforniis (the most common species in our
collection ) , males outnumbering females by
5:1. In only the largest specimens could go-
nads be recognized.

A new genus, Pseiidotindaria (not in-
cluded in the family Tindariidae), is de-
scibed. This includes species with shell
morphologies very similar to those of Tin-
daria but with soft parts that are mark-
edly different. The taxonomic affinities of
Pseiidotindaria will be discussed in a later
paper.

Pscudotindaria is siphonate. The hind
gut is arranged in a complex series of loops
and coils on either side of the body and
does not penetrate the mantle. Palp ridges
are numerous. In the case of P. erehus, the
most common species in ovn* samples, more
than 40 per cent of the specimens ha\e
recognizable gonads and the sex ratio is
even.

Shell morphologies indicate that Tindaria
and Pseiidotindaria could well be consid-
ered as recent descendants of the Paleozoic
ctenodont Protobranchia.

REFERENCES

Allex, J. A. 1971. Evolution and functional
niorphologN' of the deep water protobranch
lM\al\es of the Atlantic. Proc. Joint Occanogr.
Assembly (Tokyo 1970). 251-253.

Deep Sea Protobranchs • Sanders 6- Allen 59

\.LLEN, J. A. 1973. The adaptations of the bi-
valves of the Atlantic ab>ssal phiin. Pioc.
Challenger Soc. 4 (in press).
\llen, J. A. AND H. L. Sanders. 1966. Adapta-
tions to ab\ssal Hfe as shown by the bivalve,
Abra profuudonnn (Smith). Deep-Sea Res.,
13: 1175-1184.

\llex, J. A. AND H. L. Sanders. 1969. Nucin-
ella scrrei Lamy (Bivalvia: Piotobranchia ) ,
a monomyarian solemyid and possible living
actinodont. Malacologia, 7: 381-396.

\llen, J. A. AND H. L. San-ders. 1973. Studies
on deep sea Protobranchia. The families
Siliculidae and Lametilidae. Bull. Mus. Comp.
Zool. Harv., 145: 263-310.

\llen, J. A. AND J. F. Turner. 1974. On the
functional moiphology of the family Verticor-
diidae (Bivahia) witli descriptions of new
species from the abvssal Atlantic. Phil. Trans.
Roy. Soc. B. 268: 401-536.
LARKE, A. H., Jr. 1959. New abyssal molluscs
from off BeiTnuda collected by the Lamont
Geological Obser\atorv. Proc. Malac. Soc.
Lond., 33: 231-238.

Clarke, A. H., Jr. 1961. Ab>-ssal mollusks from
the South Atlantic Ocean. Bull. Mus. Comp.
Zool Harv., 125: 345-387.

Clarke, A. H., Jr. 1962. Annotated list and bib-
liography of the abyssal marine molluscs of the
world. Bull Nat. Mus. Can., 181: 114 pp.

Dall, W. H. 1881. Reports of the results of
dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico, and in the
Caribbean Sea, 1877-79, by the U.S. Coast
Survey Steamer 'Blake'. XV. Preliminary
report of the Mollusca. Bull. Mus. Comp.
Zool. Harv. 9: 3.3-144.

Dall, W. H. 1886. Report on the Mollusca Part
1. Brachiopoda and Pelecypoda. Reports on
the results of dredging, imder the supenision
of Alexander Agassiz, in the Gulf of Mexico
(1877-78) and in the Caribbean Sea (1879-
80) by the U.S. Coast Suney Steamer 'Blake'.
Bidl Mus. Comp. Zool Harv., 12: 171-318.

Dall, W. H. 1889. A preliminary catalogue of
the shell bearing marine mollusks and Brach-
iopoda of tlie south eastern coast of the United
States, with illustrations of manv species. Bull
U.S. Nat. Mus. 37: 1-221.

Dall, W. H. 1895. Contributions to the Ter-
tiary fauna of Florida, \\ ith special reference

to the Miocene silex-beds of Tampa and the
Pliocene beds of the Caloosahatchie River.
Tertiary mollusks of Florida Pt. III. A new
classification of the Pelecypoda. Trans. Wag-
ner Free Inst. Sci. 3: 485-570.

Kxudsex, J. 1970. The systematics and biology
of abyssal and hadal Bivalvia. Galathea Rep.
11: 7-241.

PojETA, J. 1971. A review of Ordovician pelecy-
pods. Geol Surv. Professional Paper 695:
46 pp.

Sanders, H. L. and J. A. Allen. 1973. Studies
on deep sea Protobranchia ( Bivalvia ) ; pro-
logue and the Pristiglomidae. Bidl Mus.
Comp. Zool, 145: 237-262.

ScHELTEMA, R. 1972. Reproduction and disper-
sal of bottom dwelling deep-sea invertebrates:
A speculative summary. Barobiol. & Exper.
Biol. Deep Sea: 58-66.

Smith, E. A. 1885. Report on the Lamelli-
branchia collected by H.M.S. 'Challenger'
during the years 1873-76. Rep. Scient. Res.
Challenger, 13: 341 pp.

Thiele, J. 1935. Handhuck der Systcmatischen
WcicJitierkunde, 2 Classis Bivalvia (Jena) pp.
779-1154.

Turekiax, K. K., J. K. CocHR.\N, D. p. Kharkar,
R. M. Cerrato, J. R. Vaisnys, H. L. Saxders,
J. F. Grassle ant) J. A. Allen. 1975. The
slow growth rate of a deep-sea clam deter-
mined b\- 228 Ra chronologv. Proc. Nat. Acad.
Sci. 72:' 2829-2832.

Verrill, a. E. and K. J. Bush. 1897. Revision
of the genera of Ledidae and Nuculidae of the
Atlantic coast of the United States. Amer. J.
Sci. 3: 51-63.

Verrill, A. E., axd K. J. Bush. 1898. Revision
of the deep-water Mollusca of the Atlantic
coast of North America with descriptions of
new genera and species. Part 1. Bivalvia.
Proc. U.S. Nat. Mus. 20: 777-901.

\'okes, H. E. 1967. Genera of the Bivalvia. A
systematic and bibliographic catalogue. Bull.
Am. Palcont. 51: 111-394.

YoxGE, C. M. 1939. The protobranchiate Mol-
lusca; a fvmction interpretation of their struc-
ture and evolution. Phil Trans. Roy. Soc. B.
230: 79-147.

YoNGE, C. M. 1959. The status of the Proto-
branchia in the bi\ alve Mollusca. Proc. Malac.
Soc. Lond. 33: 210-214.

■k

> .■

us ISSN 0027-410G

Sulietln OF THE

seum o
Comparative
Zoology

The American Orb-weaver Genera Cyclosa,

Metazygia and Eustala North of Mexico

(Araneae, Araneidae)

HERBERT W. LEVI

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 3
16 JUNE 1977

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Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 02138, U.S.A.

THE AMERICAN ORB-WEAVER GENERA CYCLOSA,
METAZYGIA AND EUSTALA NORTH OF MEXICO
(ARANEAE, ARANEIDAE)

HERBERT W. LEVI^

Abstract. Five species of Cijclosa, three of
Meiazygia and thirteen of Eusiala are found in the
region. One species of Cijclosa is holarctic in dis-
tribution, others are temperate and tropical Amer-
ican. The rarity of the dwarf males of the tropical
Florida Cijclosa hifurca suggests that the species
may be parthenogenetic. Metaztjgia and Ettstala
are known from the Americas only, most species
being tropical. The five temperate species of Eiis-
tala, especially the three eastern ones, are difficult
to separate; possibly they hybridize in some areas.
Two of the Eustala species are new, with the range
of southern Florida and the West Indies.

INTRODUCTION

As \\dth most orb-weaver genera, Cijclosa,
Mefozygia and Eustala have never been re-
vised and until now only some common spe-
cies could be determined with certainty.

A revisionary study such as this should
report the results of the research; that is, it
should summarize the diagnostic characters
of the species and genera revised, indicate
how to separate the species, and provide
some general information on the natural his-
toiy of the species studied.

Much previously unpublished data on
natural history can be gleaned from collect-
ing labels; the author's own experience and
published literature (if the determinations
are reliable) can supply more. A summary
of this information is of as much general
interest as are the keys and diagnosis.

On the other hand, detailed nondiagnostic

^ Museum of Comparative Zoology, Hanard
University 02138.

morphological descriptions are of little in-
terest, although they are frequently given
in revisionary studies. Of still less interest,
except to the writer, is the nomenclatural
confusion that preceded the revision. In
non-numerical, taxonomic research only the
results, not the procedures, are usually
gi\en. If the specimens key out and the
illustrations are useable, the study is demon-
strated to be adequate. Nevertheless, in
this paper I have indicated the procedures
used to study Eustala as a partial answer to
those who claim that taxonomic work might
be hastened.

ACKNOWLEDGMENTS

1 would like to thank the following per-
sons for helping in these re\'isions. W. J.
Gertsch generously made part of his un-
published manuscript on West Indian Eus-
tala a\'ailable to me. Two of the new spe-
cies from the West Indies are described
here as they also occur in southern Florida;
Gertsch's manuscript names were adopted
to avoid confusion in already labeled speci-
mens. N. I. Platnick and F. R. Wanless
went out of their way to find misplaced
specimens in their collections. R. E. Bus-
kirk, J. E. Carico, H. K. \^'allace, W. Sedg-
wick, and M. Stowe reported obsenations.
^^^ G. Eberhard, Y. D. Lubin, W. L. Brown,
A. Moreton, R. E. Buskirk, V. Brach and J.
E. Carico provided photographs. Speci-
mens were loaned bv P. H. Amaud and R.

Bull. Nkis. Comp. Zool., 148(3): 61-127, June, 1977

Bulletin Miiscitiu of Comparative Zoology, Vol. 148, No. 3

X. Schick, California Academy of Sciences;
J. A. Beatty {Cyclosa only); D. Bixler; The
British Columbia Provincial Museum, Vic-
toria; J. E. Carico; R. Crawford; C. D.
Dondalc, Canadian National Collections,
Ottawa; H. Dybas and J. B. Kethley, Field
Museum of Natural History, Chicago; W.
C. Eberhard; S. Frommer; W. J. Gertsch;
M. Grasshoff, Senckenberg Museum, Frank-
furt; M. Hubert, Museum National d'His-
toire Naturelle, Paris; B. J. Kaston; R. E.
Leech; \V. R. Icenogle; W. \V. Moss, Acad-
emy of Natural Sciences, Philadelphia; Mr.
and Ms. J. Mui-phy; W. B. Peck, Exline-Peck
Collection, Warrensburg, Missouri; N. I.
Platnick, American Museum of Natural His-
tory and Cornell University collections; S.
E. Riechert, University of Wisconsin; W. T.
Sedgwick; W. A. Shear; M. Stowe; K. J.
Stone; H. K. Wallace; C. A. Triplehorn and
A. J. Penniman, The Ohio State University
collections; F. R. Wanless, British Museum
(Natural History), London; H. V. Weems,
Florida Collection of Artlu-opods, and B. R.
Vogel. E. Mayr made comments and sugges-
tions for the introduction. Some outline
maps were suppHed by D. Quintero, L. Roth
mapped tlie species, and D. Randolph typed
numerous manuscript drafts and the final
copy. L. R. Levi corrected the syntax. The
study and its publication were supported in
part by National Science Foundation grant
BMS 75-05719. A grant from The Center
for Field Research and Earthwatch Inc.
made a trip and stay at the Archbold Bio-
logical Station, Lake Placid, Florida pos-
sible. K. Harris and J. Maluda, participants
in the field work, helped with observations
of Eustala anastem, Metazygia ivittfeldae
and some Cyclosa species.

Cyclosa, Metazygia and Eustala

Cyclosa, like Mecynogea and Cyrtophora
among the araneid orb-weavers, hangs its

Plate 1. Cyclosa conica penultimate female and a
web built by a penultimate female, New Hampshire.

eggs on a radius of the web, perhaps as a
camouflage device (Plates 1, 2). Juveniles
make a line of debris. But Cyclosa remakes
its web almost daily, as do most members of
the family, while Mecynogea and Cyrto-
phora do not. Cyclosa renews the viscid

Plate 2. Cyclosa turbinata female and her web. Upper photographs Virginia, lower one California (upper
left photo J. Carico, upper right A. Moreton, lower B. Opell).

Cyclosa, Metazygia and Eustala • Levi 63

64 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Plate 3. Cyclosa caroli. upper and middle photograph
web, bottom detail with spider (arrow) in center of
line of debris. Upper photograph south central Flor-
ida, middle and bottom Panama Canal Zone (upper
photograph J. Maluda, middle one W. Eberhard, bot-
tom Y. Lubin).

threads, leaving the egg-sacs hanging (Y.
Lubin, personal communication, Plate 4).
The holarctic Cyclosa conica is the excep-
tion. While it does hang debris and silk in
the web, it places its egg-sacs on leaves,
probably because of tlie short season in the
northern parts of the range. Uloborus, a
cribellatc orb-weaver, also hangs its egg-
sacs in the web. But cribellate silk owes its
stickiness to its woolly nature, so the webs
do not dry and have to be replaced. Fe-
males of both Uloborus and Cyclosa hang
among their egg-sacs, head up in some Cy-
closa species, and resemble their egg-sacs so
closely as to be hard to find (Plates 1-5).

Cyclosa bifurca is the only colored spe-
cies; both spider and egg-sac are green.
The female genitalia are reduced secondar-
ily, reverting almost to a haplogyne condi-
tion: there is no scape and no xentral open-
ing. Of about 350 specimens examined,
only two males were found. Is the species
parthenogenetic? The male is dwarfed and
the palpal structures are somewhat reduced.
For instance, the paramedian apophysis is
lacking and the conductor is small (Figs.
86, 87^

The accumulated errors in the literature
of several generations posed several riddles.
For instance, there has been much specula-
tion as to how the "American" Cyclosa
ociilata, common in the Mediterranean area,
was introduced to Europe (Lutz, 1915,
Simon, 1928). But Cyclosa oculata (Figs.
21-23) is actually a European species which
lias never been found in America. Because
its abdomen resembles that of the American
C. icalckcnaeri (Plate 4), Simon (1900)
confused and synonymized the two, leading
later authors astray.

Besides the poorly known species from
the Balkans, five species of Cyclosa are
known from western and southern Europe
(Roewer, 1942, Bonnet, 1956) (Figs. 21-
37 ) . Three of these are Mediterranean ( C.
alii^erica, C. sierrae and C. insulana). [C. in-
sulana is found from France and Africa to
India and the southwestern Pacific ( Bonnet,

Cyclosa, Metazygia and Eustala • Levi

I
O)

O)
CD

3
T3
CO

0)
0)

E
o

c
o

>- tn

CD u

Q. CO

- Oi

W CD

CD CO

CD .
Q.

E
o
u

(D
CD
C
<D

— 1^

CD 5
CO O

o —

>v (D

^ oJ

. E

'i' CO
(D T3

5 E

Q. O

66 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Plate 5. Cyclosa bifurca web with female and egg-sacs, 15 cm diameter, Florida (photo V. Brach).

1956).] All five species are closer to C.
conica than to the other American species.

Metazijgia is mostly made up of tropical
American species. The orbs are loose with
widely spaced spirals (Plate 6). They are
usually left up during the day, while the
spider rests in a retreat, and are replaced
every evening after dark. Metazijgia witt-
feldae, which often makes its webs on
])ridges or buildings, occupies a niche simi-
lar to that of the more nortliern Nuctenea
conuita (Clerck), and is similar in appear-
ance ( Plate 6 ) .

Eustala, although common, is not well-
known. Various species are found resting
on dead twigs of shrubs or trees. W. Eber-
hard (in letter) writes that some Eustala
have their webs up during the day, but most
(in southern Colombia) put them up in the

evening and tear them down in the morn-
ing. The webs are characteristic with some
variation. In constiiiction they are more or
less vertical and somewhat asymmetrical
with the larger part usually below the hub.
They have frame threads that do not span
paiticularly large spaces, and a hub with
several well-ordered loops and a medium
hole in the center. They are often built in
dead branches or tree tiainks. In general
they are undistinguished webs with nothing
particularly remarkable about them. My
own observations agree with Eberhard's.
Eustala anastera in central Florida make
their webs in the evening after dark. Usu-
ally the webs have disappeared by morning,
but once in awhile a web is kept ( Plate 7 ) .
Eustala and Metazijgia webs are similar
and may be horizontal or vertical. Both are

Cyclosa, Metazygia and Eustala • Levi 67

•late 6. Metazygia wittfeldae. upper left female; upper right web 15 cm horizontal diameter; lower left 18
cm horizontal diameter; lower right web with dew, 25 cm horizontal diameter.

loose constructions with few threads. And
both EmtaJa anastem and Metazygia w'ltt-
feldae are less likely than many other noc-
iturnal orb- weavers to tear down tlie web
when disturbed by artificial light or when
the web is dusted with cornstarch to make
it more visible in photographs.

METHODS

At the start of a revisionary study tlie
taxonomist has in front of him perhaps hun-
dreds of specimen collections. Are those
collected together all the same species? Can
species be separated readily by their geni-
talia, or by their size, coloration, eye ar-

68 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Plate 7. Eustala anastera, Florida; top row female; bottom webs: left with spider in web 13 cm diameter; right
spider removed, 38 cm diameter.

rangement, or the shape of the abdomen?
Perhaps a system could be based on each
character hke the one devised by Adanson,
in tlie ISth centmy, but it miglit be unnatu-
ral, each character giving a separate classifi-
cation. Some species are so distinct that the
diagnostic characters are obvious, but more
often the taxonomist has to sort out speci-
mens and try various combinations of char-
acters. Do all those that lack a hump on the
abdomen also have distinct <j:enitalic char-

acters, and do all these fall witliin a certain
size range? Might the smaller size, larger
eyes, and lack of hump reflect merely fewer
instars passed by a spider before maturity,
or do they reflect a segregated breeding
population for which we can predict also
different behavior and habits? In both
CycJosa and Eustala the numerous genitalic
differences between specimens usually rep-
resent individual variation of no taxonomic
importance. To be diagnostic, differences

Cyclosa, Metazygia and Eustala • Levi 69

must characterize all members of the popu- visers used the smallest possible sample

'ation. from a population, any specimen tliat dif-

The separation of populations from the fered would be described as new, and "dif-
mass of specimens cannot be done by intui- ficult" specimens were ignored or discarded
tion. Numerical and statistical methods are as abnormal. But these "difficult" speci-
not applical)le, as spiders grow allometric- mens represent the variation that makes re-
ally and mature after a variable number of visions challenging. The huge numbers of
instars. Statistically significant measure- specimens in American collections embrace
nients would make specimens matiu-ing in an enormous amount of variation, aside
the 8th instar distinct from those maturing from abnormalities, making a sound basis
in the 9th. This is very different among for revisionary studies,
most mites, which have a fixed number of The EusfaJa species are difficult. Cham-
molts. My own method is to find the diag- berhn and Ivie (1935) considered the
nostic characters for the population that is California-Arizona populations distinct be-
n;ost distinct, and try to delineate the more cause of a series of humps on the posterior
difficult species on the basis of the same of the abdomen, selected a juvenile speci-
characters. But is it valid to assume that men as type, and named the species rosae.
other members of the genus differ in the In 1944 in the controversial paper on the
corresponding characters? In the Cyclosa spiders of the Georgia region, the same
conica group of species (Figs. 1-37), the authors listed three soudieasteni species,
median apophysis of the palpus is a useful using (wisely in this instance) old Walcken-
character for classifying males, and differ- arian names. Eustala anastera was charac-
ences in its shape correlate witli differences terized as having "a shai"p conical tip to the
in size, shape of abdomen, and other char- abdomen. It occurs in a great variety of
acters. The shape of the median apophysis color patterns. . . ." Nothing was said about
is similarly useful in other American species how to separate males. "Eustala cepina is
of Cyclosa, but for separating species of smaller than anastera, the abdomen is less
Lariiiia and Eustala, it is useless. To find sharply angulate and is broad. It lacks the
the most useful characters, I make numer- silky white hairs fovmd on the top of the
ous outline drawings to scale, few of which head of anastera and triflex, or at most are
will be used in the final presentation of the much reduced." Eustala triflex ( = enier-
research. The nontaxonomist, who sees toni) "differs from anastera and cepina
only conclusions, remains puzzled as to the which occur in the same region by larking
aims and methods of the study. [sic] the terminal angulation on the abdo-

In Cyclosa the females of North American men, the abdomen being rounded behind,
species could readily be separated by the There are also differences in the male pal-
shape of the abdomen as well as by the pus." But what are these differences in the
epigynum. Males of one species were all complicated palpus?

accompanied by females, and all came from It is not surprising that Kaston (1948) in

the northern part of North America. But his discussion on Eustala anastera, does not

males of other species were much more dif- list cepina and triflex in the synonymy, but

ficult to separate. An occasional specimen indicates that "Chamberlin and Ivie prefer

matched females or accompanied females, to maintain [triflex] as a distinct species "

At first the palpi could not readily be sepa- Chickering (1955), describing Central Amer-

rated, but gradually differences were found ican Eustala, listed cepina as a synonym of

in the median apophysis (Figs. 47, 60, 73). anastera, hnt did not mention triflex. Archer

Today's approach is vastly different from ( 1951 ) , however, considered triflex distinct

that of twenty to fifty years ago, when re- and described a fourth species, E. arkansana,

70 Bulletin Muscitiu of Comparative Zoology, Vol. 148, No. 3

citing differences of the epigyna (wliich
I could not \erify). For arkamana, Archer
ilhistrated the epigynum, and for all the
species he illustrated a palpal stmcture he
called the "median apophysis." I cannot rec-
ognize the palpal structure illustrated; it
may be the embolus. Archer did not indicate
whether he illustrated the one from the left
or the right palpus.

I started my investigation of Eustola with
the American Museum collection. Gertsch,
Archer and hie had used the Chamberlin
and I\'ie ( 1944 ) names on only a few spec-
imens (most others were not determined),
but Chickering, who examined a small part
of the collection, labeled all specimens E.
anmtera. One locality, in eastern Pennsyl-
\'ania, yielded a large series of males and
females. These had labels of the three spe-
cies {anmtera, cepina, "triflex"), and I
started to draw tliese and study their differ-
ences. But I could not find the differences
when I tried to use tliem for separating
other collections. Only one character, the
ventral row of macrosetae on the second
femur of E. anastera, remained constant
(Fig. 214), and I subsequently sorted out
all collections using these "spines." How-
ever, such large setae could reflect allo-
metric growth, as the anastera were also the
largest specimens. I removed all E. rosae
because they seemed distinct and similar
in size to anastera, lacked the femoral
macrosetae in males, had humps on the ab-
domen, and had a distinctive epigynum
(Fig. 193). To get some new ideas, I ex-
amined E. californicnsis, a distinct North
American species. The diagnostic genitalic
characters were a differently shaped termi-
nal apophysis and median apophysis in the
male palpus (Figs. 147, 148), and the shape
of the posterolateral plates of the epigynum
(Fig. 139). The embolus of the palpus,
which I had previously carefully examined
and illusbated was not significantly differ-
ent. Because it is soft, the median apophysis
is a more difficult character to work with.
With this new knowledge I returned to E.

rosea and confirmed that all females with'
extra abdominal humps (Fig. 196) also had |
the extra dorsal lobe on the posterior face of ||
the epigynum (Fig. 193). The males had
very distinct "half-spear-shaped" terminal
apophyses (Figs. 202, 203, 313) and lacked
the ventral setae on the second femur ( Fig.
201), characteristic of anastera. I returned
to anastera only to find that it had a similar
but less distinct "half-spear-shaped" termi-
nal apophysis (Figs. 215, 216). One speci-
men, determined by Ivie to be anastera, was
an exception (Fig. 315). (It turned out to
be one of many males with this character.)

Could the bubble-shaped transparent
subterminal apophysis (sa in Fig. 232),
which differed also in E. caViforniensh, be
used? Sketches and comparisons indicated
not. But the combined characters of termi-
nal apophysis shape, femoral setae, large
size and heavy pigmentation, facilitated
sorting out anastera. Only one specimen
with an anastera palp lacked femoral setae.
And, several specimens had filamentous
terminal apophyses, but were large, dark,
and had femoral setae. Do these excep-
tional specimens represent a new species,
intermediates, or perhaps products of intro-
gression? Adopting inti-ogression as a tem-
porary working hypothesis, I began to
doubt that I could separate most males of
this species. The disturbing thought oc-
curred to me that Ivie, in working tlir-ough
the three large collections from one locality
in Pennsylvania, might not have kept diffi-
cult-to-place intennediate specimens with
the labeled collections.

I decided to re-examine "triflex" males.
Preliminary examination had shown the
palpus to have an especially long tail on the
conductor (Figs. 266, 309-311). Careful
examination indicated that the long tail
usually was associated with a relatively
short terminal apophysis (Figs. 309-311).
But there were specimens with a half-spear-
shaped embolus. Was triflex merely a small
E. anastera that failed to grow the femoral
macrosetae? Some specimens had the char-

Cyclosa, Metazycia and Eustala • Levi 71

acteristic long tail of the conductor but had
a long terminal apophysis, as does A. cepina.
Are these separate species or hybrids? The
females associated with these males had the
oval abdomen (Fig. 258) described by
Chamberlin and Ivie, and also had an epig-
ynum distinct from that of anastera, but
apparently not from cepina (as far as I
could tell at this point). My confidence in-
creased and I decided to work on a new,
\'ery distinct species found in the West In-
dies and Florida (E. cazieri) to find some
new characters to use before returning to
the male E. anmtera for measurements and
recording collecting data.

Examination of the new species provided
e\'idence tliat the main pattern of macro-
setae is not merely the result of allometric
growth: tire large E. rosae has no macro-
setae ventrally on the second femur (Fig.
201), the small E. cazieri has manv (Fig.
il35).

The next problem was to separate females
of E. cepina. By making numerous draw-
ings and clearing epigyna, I fomid that in
cepina the base of the epigynum has a tiny
sclerotized scale dorsally (Figs. 286-290).
The scale is absent in E. "triflex" and E.
anmtera, and absent or transparent in Flor-
ida specimens believed to belong to the spe-
cies. This character was abandoned late in
the study as it is not consistent.

To make sure that these Florida females,
which are much smaller than E. anastera
from the rest of the range, really are the
same species, I decided to study the asso-
ciated males. I went back to various other
West Indian species, and found that the
males of some differed in the shape of the
conductor. I now made outline drawings
of the conductor, only to discover that the
long tail of E. "triflex" was more distinct
than I previously thought and, more im-
portantly, tliat of these three species, E.
cepina lacked the distal ventral lobe of the
conductor ( upper right in Figures 303-308 ) ,
proxiding an additional diagnostic charac-
ter. It also reaffirmed the differences pre-

viously noted and compoTindcd the problem
that E. anastera comes with short (Fig. 315)
and long (Fig. 314) teraiinal apophyses. It
seemed that if two species are collected to-
gether, specimens are easy to separate, but
lone individuals often cannot be placed.

In consulting type specimens and Abbot
illusti'ations, I found that die illustrations
labeled Epeira triflex were contrastingly
marked (Figs. 219, 222), a coloration found
only in E. anastera, not in any specimen I
had available of E. ''triflex." The name was
changed to E. emertoni, the next oldest
name available.

After the first triumph of finally figuring
out that there are differences and the spe-
cies can be told apart, there comes doubt
about being able to place all specimens. If
all specimens can be determined, new
doubts arise: now that I have finally dis-
covered the obvious, is it worth publishing?
But after the doubts are overcome, there
comes the challenge to put the conclusions
into a fonn that a nonspecialist can use for
separating species. The last generation's
specialists indulged in a kind of games-
manship, just stating that there are differ-
ences, but failing to describe them, or hid-
ing them in drawings that made comparison
impossible.

After examining, measuring and deter-
mining the contents of a few hrmdred vials,
most with many specimens, I found deter-
minations getting easier. I had uncon-
sciously used the relative size of the conduc-
tor in males and found now that not only
does E. "triflex" (= emertoni) have a rela-
tively large conductor (Figs. 309-311),
Init also that anastera has a relatively small
conductor (Figs. 298-302).

While determining specimens and record-
ing data, I took measurements of specimens
from all parts of the range and found that
the senitalia of anastera and the similar ce-
pina do not overlap in size (Figs. 283-290
and 298-308). That the area of tlie conduc-
tor "above" the embolus is larger in anastera
than in cepina and emertoni, I did not re-

72 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

alize until linishing the illustraUons (Figs. Remaining questiom. Other questions

298-311). remain um-esolved. Is Eiistala conchlea

But a lew problem specimens remained: (Figs. 269-279) just a western forai of £.

the male cinmtera with tlie terminal apophy- emertoni with a large abdominal hump? It

sis short (Fig. 315), and those lacking was arbitrarily decided to keep them sepa-

macrosetae on the second femur; a few rate.

emertoni {= "triflex') with the terminal Among tlie puzzHng specimens are the

apophysis long; and a few cepina with a asymmetrical ones. The macrosetae on left

distal lobe on the conductor. Problem fe- and right femora of the same specimens

males may have been overlooked due to often differ, or there may be one ventral

relati\ely arbitraiy decisions in separating macroseta on one second femur, none on the

specimens with fewer characters. other. (In the E. amstera group of species

At the end, when determining various these were all considered £. arw^stera.) An

borrowed collections, I found that Eustala interesting epigynum is that of an E. cepina

anastcra from Nova Scotia and some from female from Emmet County, Michigan (A.

Quebec have smaller abdominal humps, M. Chickering, collector, in the Museum of

males have a much smaller conductor ( Fig. Comparative Zoology) (Fig. 248). Only H

298), and females have the middle piece of the epigynum is asymmetrical, not tlie rest

the epigynum larger (Fig. 229). Are they of the animal. More startling is the left pal-

a distinct species? One male from Ontario pus of a male E. anastera from Kisatchie

had one Nova Scotia-like palpus and one National Forest, Grant Parish, Louisiana

"normal" anustcra palpus. I decided that (A. F. Archer, collector, in tlie American

perhaps one palpus was more dehydrated Museum of Natural History). The left pal-

and shri\ ellcd. pus has a unique bulbous terminal apophy-

A single collection, from Jefferson Co. in sis, the right one a normal, short, pointed

northern Florida, included botli small fe- one (Fig. 231).

males of the central Florida type, and larger Several collections of the Eustala anastera

sized females similarly marked. Are the group might be new species, but I am hesi-

smaller ones a distinct species? Adult fe- tant to name them until more specimens are

males collected at Archbold Biological Sta- available. One collection, consisting of two

tion in Febmary and March 1976 were all females and a male that appears to be a

considerably larger than females from the small E. anastera, conies from Black Mesa

same population collected in July and State Park, Cimarron County, Oklahoma

August 1975 by M. Stowe. The winter ones (16 August 1964, H. Fitch in tlie American

were predominantly brown, the summer Museum of Natural History). The females

ones green on the abdomen. have three posterior humps in a row on the

Character displacement. Eastern Cana- abdomen, and the epigynum in posterior
dian specimens of E. rtWisf era are of slightly view has a very small middle piece (Fig.
smaller size — the males ha\ing few macro- 230). The male lacks macrosetae on the
setae and a smaller conductor (Fig. 298) venter of the second femur; the conductor
and the females having a smaller hump and of the palpus is like that of E. cepina, and
larger epigynal middle piece (Fig. 229)— lacks the large lateral conductor lobe pres-
resenible E. cepina and E. emertoni, and are ent in most specimens of E. ana.^era.
more distinct in the shared range of all three Another new species may be represented
species because of character displacement, by two females, one from Cologne (? Go-
Perhaps the Mexican specimens of this Had Co.), Texas (7 June 1937, S. Mulaik in
group all belong to E. anastera but look dif- the American Museum of Natrn-al History)
ferent in the absence of competing species, and another from Little Pine Key, Florida

Cyclosa, Metazygia and Eustala • Levi 73

(27 March 1939 in the American Museum
of Natiual History). Both look hke very
large E. anastera with the more common
spotted patchy pattern, but more contrast-
ing. The females have two posterior humps
and, in the epigynum, a very heavy, large
scape with parallel sides (Fig. 228). The
Texas female, whose epigynum was illus-
trated (Fig. 228), was 14.0 mm in total
length, carapace 4.9 mm long, 4.1 wide.
Several very large E. anastera males from
southern Florida (as compared with the
small E. anastera from central Florida) may
belong with these females. But the question
remains whether they represent a separate
species or large-sized populations.

After all was completed, the "easy" Meta-
zygia illustrated and determined, I felt that
I had to return to these difficult Eustala
specimens. Perhaps I had overlooked char-
acters seen in ventral view of the median
apophysis, the paracymbium and sculptur-
ing of the tegulum of the palpus. But no
new characters were found. However, on
reexamination, the Oklahoma male turned
out to be E. cepina, the females (Fig. 230)
are perhaps a new species to be named when
additional specimens are available. I also
reexamined most Texas and southern Flor-
ida specimens of E. anastera in search for
additional large females having an epigynal
scape with parallel sides (Fig. 228), and
perhaps for males, but found only intermedi-
ates, all in the collection from Raven Ranch,
Kerr County, Texas. The intermediate spec-
imens (Figs. 226, 227) dissuaded me from
describing the two specimens as a new spe-
cies.

While revision of Eustala is now com-
pleted, I would not be suiprised to find
additional sibling species among the Eustala
anastera collections.

Cyclosa Menge

Cyclosa Menge, 1866, Schrift. naturforsch. Ge-
sellsch. Danzig, neue Folge, 1 : 73. Type species
C. conica (Pallas) by nionotypy. The name is
feminine.

Farazijgia di Caporiacco, 1955, Acta biol. Vene-

zuelica, 1: 345. Type species P. accentonotata
di Caporiacco [=C. caroli (Hentz)] by mono-
typy. NEW SYNONYMY.

Diagnosis. Cyclosa species differ from
those of other Araneidae genera and espe-
cially from Araneus in the narrow head re-
gion of the carapace, often separated by
shallow grooves from the thoracic region
(Figs. 10, 12, 29, 48). The eyes are closely
spaced, posterior median eyes almost touch-
ing (Figs. 10, 12, 14). Cyclosa differs from
Larinia, which also has the posterior median
e>'es close, in having banded legs, and in the
shape and coloration of the abdomen. There
are dorsal, paired, black or gray patches on
white (Figs. 10, 29, 48, 61, 74) and a char-
acteristic pair of ventral white spots sur-
rounded and separated by a black band
running from epigynum to spinnerets and
posteriorly surrounding the spinnerets (Figs.
11, 49, 62, 75). The posterior dorsal end of
the abdomen is extended beyond the spin-
nerets in the female and there may be shoul-
der humps or additional posterior humps
( Figs. 2, 10, 28, 29, 39, 48, 52, 61, 65, 74, 78,
88). Cyclosa further differs from Araneus
and Larinia in that the male palpal patella
has only one macroseta (Fig. 1).

The web is diurnal, its form diagnostic;
lightly spun with few frame threads, it has
a stabilimentum containing debris or a verti-
cal row of egg-sacs through the center; the
spider rests at the lower end or in a gap in
the decoration and is often difficult to find
(Plates 1-5).

Description. The head region of the
brown carapace is narrow and lighter in
color than the thoracic region; the thoracic
depression is round (Figs. 10, 29, 48, 61,
74). The carapace is covered with down.
The anterior median eyes are slighth' larger
than the others, which are subequal in size
(Fig. 16). The anterior median eyes are
their diameter apart, usually one, but not
more than two and one-half diameters from
laterals. Posterior median eyes touching or
less than their diameter apart, one and one-
half to three diameters from laterals ( Figs.
10, 29, 48, 61, 74, 88). The clypeus height

74 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

equals about the diameter of the anterior
median eyes (Fig. 16). The sternum is dark
brown, often enclosing white pigment
patches. The coxae are light, sometimes
with dark marks. The legs are light with
dark bands with short setae and macrosetae.
The spinnerets are usually dark brown.
Cijclosa bifurca departs from the drab color-
ation of other species by being green. The
widespread C. imulana (Fig. 29) of Eurasia
and Africa to the Pacific has a silvery abdo-
men, perhaps an adaptation to the open
sunny areas it frequents (M. H. Robinson
et al, 1974).

Males are smaller than females, more
sclerotized, darker in color, and have the
abdomen almost spherical with humps only
faintly indicated. The markings are dark
with few paired light spots, but with some
indications of the humps (Figs. 1, 12, 38,
50, 51, 63, 64, 76, 89). The endites have a
lateral tubercle facing a minute cone on the
palpal femur (lacking in the small male of
C. bifurca). The first coxa of the male has
a small hook (also lacking in C. bifurca)
that fits into a groove of the male second
femur. In addition, the fourth coxae of C.
conica are anned with two macrosetae (Fig.
15). The second tibia is only slightly
thicker than the first with a few more
macrosetae (Figs. 1, 12, 50, 51, 63). The
male of C. bifurca is dwarfed (Fig. 89).

Genitalia. The epigynum has a small weak
scape, the shape of which may be diagnos-
tic: straight and pointed in C. conica (Fig.
4), oval in C. iurbinata (Fig. 41), almost
circular in C. caroli (Fig. 54), and usually
with parallel sides in C. walckenaeri (Fig.
67). The scape is absent in C. bifurca (Fig.
80). The seminal receptacles of all are
sclerotized (Figs. 3, 5, 40, 42), but the con-
necting ducts and their openings to the out-
side are so thin-walled that they are hard to
find, and once found their course is difficult
to follow. They open in a fold on the venter
of the base on each side of the scape (Figs.
40, 53), except that in C. bifurca they open
on the posterior not far from the fertiliza-

tion ducts (Figs. 79, 81), a peculiar, prob-
ably secondary modification approaching
haplogyne condition. Some material may
be found in the depression having the open-
ings of the epigynum, but I believe that
these are an epigynal plug formed from
mucus and not a part of the palpus left be-
hind. (It is not known whether males or
females can mate several times. )

The palpal patella has one macroseta
(Fig. 1). The bulb has a large conductor
holding the tip of the embolus ( "c" in Figs.
7, 17, 20), a small terminal apophysis ("a"
in Figs. 17, 20), and a paramedian apophy-
sis (pm), the latter apparently absent in
C. bifurca. The embolus (e) is thread-
shaped in all and the median apophysis has
moved to the ventral side in all except C
bifurca ("m" in Figs. 8, 17, 20, 46). The
complex median apophysis is species spe-
cific (Figs. 9, 23, 27, 33, 37, 47, 60, 73, 87),
apparently fitting the epigynal scape into
which it hooks during mating.

Natural History. The web has few frame
threads (Plates 1-5). That of C. conica is
almost circular, with about 40 radii (Wiehle,
1931); in each sector are 20-30 viscid
threads separated by 2 to 3 mm ( Plate 1 ) .

The spider hangs in the middle of the
web; juveniles have a detritis-covered sta-
bilimentimi (Plates 1, 4). Adults place the
egg-sacs in a vertical line in the center, the
spider resting at one end or the middle
(Plates 1-5). Only Cijclosa conica does not
place her egg-sacs in the web, no doubt an
adaptation to the short season of the more
northern areas it frequents. The sacs are
left hanging when the viscid threads of the
web are renewed (Y. Lubin, personal com-
munication).

Some Cijclosa species are known to hang
with the head up rather than down, like the
widespread Eurasian Cijclosa imulana
(Wiehle, 1928).

The shape of the stabilimentum of young
Cijclosa cannot be used to separate species
as it may differ greatly even in successive
webs of the same individual (Marson, 1947).

Cyclosa, Metazygia and Bust ALA • Le

'•?.. \'

Cyclosa conica

Map 1. Distribution of Cyclosa conica (Pallas) in North America.

All species shake the web when disturbed,
then may drop on a thread.

Species. Of the five species north of
Mexico, C. conica is northern and holarctic,
the remainder southern (Maps 1, 2). There
are numerous tropical American species.
The species north of Mexico can be sepa-
rated by the shape of the abdomen of the
female (Figs. 2, 39, 52, 65, 78), the shape of
the epigynum, especially the scape (Figs.
4, 41, 54, 67), and the matching median
apophysis of the palpus ( Figs. 9, 47, 60, 73 ) .
It is ironic that A. Archer, who tried to
separate all Araneidae species on the shape
of the median apophysis alone, did not
study Cyclosa, one genus in which the stiaic-
ture is of diagnostic importance.

Distribution. Cyclosa species are found
in all parts of the world.

Key to female Cyclosa north of Mexico

1. Posterior tip of abdomen biforked (Fig. 88);

epigynum without scape (Figs. 80, 84);

Florida, Alabama coast and southern Texas

hifurca

- Abdomen with a single posterior hump ( Fig.

10) or four posterior tubercles (Fig. 74)

2(1 J Epigynum with sclerotized lobe on each
side of scape (Fig. 4); Alaska, south to
\'irginia, Arizona and California conica

- Epigynum base without sclerotized lateral

lobes (Figs. 41, 67); Connecticut to Wash-
ington and south 3

3(2) Abdomen with a pair of dorsal tubercles
on anterior half of abdomen ( Figs. 39, 65 )
4

- Abdomen without dorsal tubercles; abdomen

posterior to spinnerets longer than part in
front of spinnerets (Fig. 62), epigynum
scape an oval to circular lobe (Fig. 54);

Georgia to Texas caroli

4(3) Abdomen with a single posterior hump
(Fig. 39), epigynal scape a rounded lobe

76 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Cyclosa walckenaeri

Cyclosa bifurca

Map 2. Distribution of Cyclosa turbinata (Walckenaer), C. caroli (Hentz), C. walckenaeri (O.P.-Cambridge)
and C. bifurca (McCook).

Cyclosa, Metazygia and Evstala • Levi 77

Figures 1-12. Cyclosa conica (Pallas). 1. Male from side. 2. Female from side. 3-6. Epigynum: 3, 4. Ven-
tral, 5, 6. Posterior. 3, 5. Cleared. 7-9. Left male palpus: 7. Mesal. 8. Ventral. 9. Median apophysis,
ventral. 10. Female, dorsal. 11. Female abdomen, ventral. 12. Male, dorsal.
Scale lines. 0.1 mm, except Figs. 1, 2, 10-12, 1.0 mm.

(Fig. 41); from Connecticut to Washing-
ton and soutli tuihinafa

Abdomen with four posterior humps (Fig.
65), epigynal scape usually with sides par-
allel ( Fig. 67 ) , southern Florida, southern
Texas, California walckenaeri

Key to male Cyclosa north of Mexico

1. Fourth coxae each with a pair of macrosetae
(Fig. 15); palpus with median apophysis
heavily sclerotized and its distal tip folded
o\er ( Fig. 9 ) ; Alaska south to \' irginia,
Arizona and California conica

78 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

- Fourth coxae ne\er with macrosetae (Fig.

77); median apophysis Ughtly sclerotized,

tip not folded 2

2(1) Median apophysis niesal (Fig. 86); tip
of abdomen biforked (Fig. 89); total
length less than 2 mm; Florida, Alabama
coast and southern Texas bifurca

- Median apophysis almost hidden in mesal

view (Figs. 45, 58, 71), posterior tip of
abdomen with a tubercle or four tubercles;

total length more than 2.1 mm 3

3(2) Abdomen usually extended beyond spin-
nerets (Fig. 51); median apophysis of the
palpus short without a middle spine but
with a rounded keel proximal to distal tip
(Fig. 60); Georgia to Texas cawli

- Abdomen with only a posterior hump, or

four slight posterior hmnps ( Figs. 38, 64 ) ;
median apophysis with a spine in middle,
with or without distal keel ( Figs. 47, 73 )

4(3) Posterior tip of abdomen usually with
indications of four tubercles ( Fig. 64 ) ;
median apophysis long, with a tiny median
spine and a more distal keel ending in
distal tip (Fig. 73); soudiem Florida,
southern Texas, California ivalckenaeri

- Posterior of abdomen with at most a dorsal

hump ( Fig. 38 ) ; median apophysis with a
large median spine but no keel distally in
ventral view (Fig. 47), in subapical view
keel e.xtending from distal to median spine,
from Connecticut to Washington and
south tiiibinata

Cyclosa conica (Pallas)

Plate 1, Figures 1-19, Map 1

Aranea conica Pallas, 1772, Spicilegia Zoologica,
9: 48, pi. 1, fig. 16. Female specimen from
Germany, believed lost.

Epeira canadensis Blackwell, 1846, Ann. Mag.
Natur. Hist. (ser. 1), 17: 81. Juvenile type
from vicinity of Toronto, in the Hope Museum
at Oxford, lost.

Cyclosa conica, — Emerton, 1884, Trans. Connecti-
cut Acad. Sci., 6: 321, pi. 34, fig. 3, pi. 38, fig.
11, 9, $. Keyserling, 1893, Spinnen Amerikas,
4: 276, pi. 14, fig. 205, 2,6- McCook, 1894,
American Spiders, 3: 225, pi. 17, figs. 3-4, $,
S . Emerton, 1902, Common Spiders, p. 183,
figs. 428, 429, 9, S. F.O.P.-Cambridge, 1904,
Biologia Centrali-Americana, Araneidea, 2: 493,
pi. 46, figs. 19, 20, 9, 6. Wiehle, 1931, in Dahl,
Tierwelt Deutschlands, 23: 18, figs. 8, 17-21,
9, S. Comstock, 1940, Spider Book, rev. ed.,
p. 465, figs. 463-464, 9, web. Roewer, 1942,
Katalog der Araneae, 1: 754. Kaston, 1948, Bull.

Connecticut Geol. Natur. Hist, 70: 236, figs.
711-713, fig. 2037, 9, S, web. Locket and
Millidge, 1953, British Spiders, 2: 166, fig. Ill,
9 , S . Bonnet, 1956, Bibliographia Araneorum,
2: 1310.

Note. Many specimens of C. turbinata in
collections had been erroneously labeled as
C. conica, thus literature citations of "C.
conica" are not reliable, and records from
the southern states, Mexico, Central and
South America are all erroneous.

Measurements. Female from Wyoming:
Total length 5.5 mm. Carapace 1.9 mm
long, 1.4 wide. First femur, 2.1 mm; patella
and tibia, 2.3; metatarsus, 1.4; tarsus, 0.7.
Second patella and tibia, 2.0 mm; third, 1.3;
fourth, 1.9.

Male from Wyoming: Total length 3.5
mm. Carapace 2.2 mm long, 1.6 wide.
Head 0.7 mm wide. First femur, 2.6 mm;
patella and tibia, 2.7; metatarsus, 1.6; tarsus,
0.7. Second patella and tibia, 2.1 mm; third,
1.4; fourth, 1.7.

Variation. Females vary in total length
3.6 to 7.9 mm, carapace 1.7 to 2.5 long, 1.3
to 1.7 wide. Males vary in total length 3.5
to 4.9 mm, carapace 2.0 to 2.3 long, 1.5 to
1.6 wide. The largest specimens came from
the northeastern states. Specimens from
Oregon and Washington had greater size
variation dian those from other parts of the
range. The caudal hump varies in length
and is quite long in some populations (Figs.
13, 14). Rarely are females almost all black.
All long-tailed and black indixiduals came
from the southernmost localities. One spec-
imen (Fig. 13) had a long tail as well as a
relatively long epigynal scape with its tip
twisted.

Diaiinosis. In North America C. conica
is the only Cyclosa species over most of its
range; only in the south does its range over-
lap with that of C. turbinata. Cyclosa
conica is larger than C turbinata and lacks
the two anterior dorsal abdominal humps
(Figs. 2, 10). The epigynum base in C
conica has a sclerotized lobe on each side of
the scape (Figs. 4, 6), unlike C. turbinata,
and the median apophysis of the palpus is

Cyclosa, Metazygia and Eustala • Levi 79

Figures 13-19. Cyclosa conica (Pallas). 13, 14. Female abdomen: 13. (Southern California) 14. (Minnesota)
15. Male, fourth coxae, ventral. 16. Eye region and chelicerae of female. 17-19. Left male palpus, expanded
(17, 19, without cymbium). 17. Submesal view. 18. Subdorsal view. 19. Embolic division, dorsal.

Figure 20. Cyclosa turbinata (Walckenaer) male palpus, expanded, submesal view.

Abbreviations, a, terminal apophysis; c, conductor; e, embolus; h, hematodocha; m. median apophysis;
pm, paramedian apophysis; r, radix; t, tegulum; y, cymbium.

Scale lines. Figs. 13-15, 1.0 mm; Figs. 17-20, 0.1 mm.

sclerotized, its distal tip folded over and
pointed (Figs. 8, 9, 17). Two macrosetae
on the fourth coxae of males (Fig. 15) are
only rarely absent. In soutliem Europe C.
conica can be confused with tlie very simi-
lar C. sierrae Simon (Figs. 30-33) and C.
algerica Simon (Figs. 34-37). The males of
these also have two macrosetae on the
fourth coxa.

Natural Histonj. The orb of C. conica is
found on shiaibs and understory of conifer-
ous forests, sometimes deciduous, where it
is the most common orb-weaver. According
to Kaston ( 1948 ) the orb is wider than liigh,

with 40 to 50 radii, and lacks a retreat, tlie
spider resting in tlie center (Plate 1). When
disturbed the spider shakes the web or may
drop out of the web. There may or may not
be a stabilimentum in webs of the same
individual. Objects falling into tlie web and
insect remains are incoi^porated into the
stabilimentum. Matiue males do not build
orbs. The three to five egg-sacs of loose
silk are elliptical, yello\vish brown, 3x7
mm, and are attached to dead twigs or
under leaves, but not to the orb. The egg-
sacs contain 10 to 130 eggs ( Kaston, 1948 ) .
Males are mature from May to July in

Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

New England and from March to June in
California. Mature females can be found
from June to late August in New England
and from March to September in California.
The species overwinters in juvenile stages.
Distribution. Holarctic, in America from
Alaska to southern West Virginia, southern
Illinois to southern New Mexico and Baja
California Norte ( Map 1 ) .

Cyclosa turbinata (Walckenaer)
Plate 2, Figures 20, 38-50, Map 2

Epeira turbinata Walckenaer, 1841, Histoire Na-
hirelle des Insectes Apteres, 2: 140. Female
types are figures no. 79 and 80 from Georgia in
Abbot's Georgia Spiders manuscript in the British
Museum, Natural History. Copy in the Museum
of Comparative Zoology, examined.^

Epeira caudata Hentz, 1850, J. Boston Soc. Natur.
Hist., 6: 23, pi. .3, fig. 14, $. Female types from
United States in Boston Natural History Mu-
seum, destroyed.

Singa vanbrmjsselii Becker, 1879, Ann. Soc. Ento-
mol. Belgique, 22: 78, pi. 1, figs. 4-6, S. Male
holotype from Pascagoula, Mississippi in the
Institut Royal des Sciences Naturelles de Belg-
ique, Brussels, examined.

Cyclosa index O.P.-Cambridge, 1889, Biologia
Centrali-Americana, Araneidea, 1: 51, pi. 6, fig.
6, 9 . Female holotype from Tamahu, Guate-
mala in the British Museum, Natural History,
examined. F.P.-Cambridge, 1904, Biologia Cen-
trali-Americana, Araneidea, 2: 496, pi. 47, fig.
12, 9 . NEW SYNONYMY.

^ Note added in proof. C. Dondale made me
aware recently that, according to Article 72 of the
International Code of Zoological Nomenclature, the
type has to be a specimen; thus the Abbot illustra-
tion cannot be the type. A neotype may be desig-
nated (Art. 75); this has not been done here.

Cyclosa caudata, — Keyserling, 1893, Spinnen Anier-
ikas, 4: 279, pi. 14, fig. 206, 9,6-

Cyclosa culta O.P.-Cambridge, 1893, Biologia Cen-
trali-Americana, 1: 112, pi. 14, fig. 12, S. Two
male syntypes from near Omilteme, Guerrero,
Mexico in the British Museum, Natural History,
examined. F.P.-Cambridge, 1904, Biologia Cen-
tiali- Americana, Araneidea, 2: 493, pi. 47, fig. 2,
c^. NEW SYNONYMY.

? Cyclosa tuberculifera O.P.-Cambridge, 1898,
Biologia Centrali-Americana, Araneidea, 1: 269,
pi. 36, fig. 10, $ . Male holotype without palpi
from Teapa, Mexico in the British Museum,
Natural History, examined. F.P.-Cambridge,
1904, Biologia Centrali-Americana, Araneidea,
2: 493, pi. 47, fig. 1, £. Doubtful NEW SYN-
ONYMY.

Cyclosa turbinata, — McCook, 1893, American Spi-
ders, 3: 224, pi. 17, figs. 5, 6, $, c^ . Comstock,
1940, Spider Book, p. 468, fig. 467, $ . Roewer,
1942, Katalog der Araneae, p. 761. Kaston, 1948,
Bull. Connecticut Geol. Natiu". Hist. Sui-v., 70:
237, fig. 710, 9. Bonnet, 1956, BibHographia
Araneorum, 2: 1325.

Cyclosa nanna Ivie and Barrows, 1935, Bull. Univ.
Utah, biol. ser. 3(2): 18, figs. 52, 53, 9, S.
Male holotype and female paratype from Naples,
Georgia, lost. NEW SYNONYMY.

Note. Specimens of C. nanna are not in
the American Museum or University of
Utah or Ohio State University collections.
The illustration shows the epigynum of C.
turbinata.

Specimens in many collections of C. tur-
binata had been misidentified as C. conica.
Many C. conica records appear to be this
species.

Measurements. Female from Louisiana:
Total length 4.3 mm. Carapace 1.5 mm long,
1.0 wide. First femur, 1.3 mm; patella and

Figures 21-37. Old-world Cyclosa.

Figures 21-23. C. oculata (Walckenaer) (Central Europe): 21, 22. Epigynum. 21. Ventral. 22. Posterior. 23.
Left male palpus, mesa! view.

Figures 24-29. C, insulana (Costa): 24-26. Epigynum: 24, 25. Ventral. 26. Posterior. 27. Palpus, mesal view.
28. Female abdomen from side. 29. Female, legs removed. 24, 26, 28, 29. (Southern France). 25, 27. (New
Guinea).

Figures 30-33. C. s/e/rae Simon (Centralltaly): 30, 31. Epigynum: 30. Ventral. 31. Posterior. 32 33. Palpus-
32. Mesal. 33. Ventral.

Figures 34-37. C. algerica Simon (Southern France): 34, 35. Epigynum: 34. Ventral. 35. Posterior. 36, 37.
Palpus: 36. Mesal. 37. Ventral.

Scale lines, 0.1 mm. Figs. 28, 29, 1 mm.

Cyclosa, Metazygia and Eustala • Levi 81

Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

tibia, 1.4; metatarsus, 0.8; tarsus, 0.5. Sec-
ond patella and tibia, 1.2 mm; third, 0.8;
fourth, 1.3.

Male from Louisiana: Total length 2.5
mm. Carapace 1.4 mm long, 1.1 wide. First
femur, 1.2 mm; patella and tibia, 1.3; meta-
tarsus, 0.7; tarsus, 0.4. Second patella and
tibia, 1.1 mm; third, 0.7; fourth, 1.0.

Variation. Females vary in total length
3.3 to 5.2 mm, carapace 1.4 to 1.7 long, 0.9
to 1.3 wide. Males vary in total length 2.1
to 3.2 mm, carapace 1.3 to 1.6 long, 1.0 to
1.2 wide. Small and large individuals ap-
peared in many collections, but Florida
specimens are usually small. Some females
from Central America have a much longer
posterior tail.

Diagnosis. Female C. turbinata differ
from C. conica by being smaller in size and
having a pair- of anterior dorsal humps
(often indistinct) on the abdomen (Figs.
39, 48) and by the lightly sclerotized base
of the epigynum (Fig. 41). In North Amer-
ica, females differ from other species by the
abdomen shape and the details of the epigy-
num (Figs. 39, 41). The males differ from
those of C. conica by lacking macrosetae on
the fourth coxae, and by their smaller size
(less than 3.3 mm total length); from C.
caroli by usually having the abdomen only
slightly overhanging the spinnerets (Fig.
38); and from the related C. caroli and C.
walckenaeri by having a median apophysis
in the palpus with a terminal hook and a
large median proximally directed tooth
(Figs. 46, 47). There is no such tooth in C.
caroli, and that of C. tvalckenaeri is small
and the median apophysis is relatively
longer.

Natural History. The web with stabili-
mentum is illustrated in Plate 2. The co-
coons are attached to the stabilimentum and
are covered with insect remains. The lowest
ones may have spiderlings wliile the upper
one has only eggs (Kaston, 1948).

Specimens have been collected by sweep-
ing lawns in West Virginia, by sweeping
abandoned fields and in a garden in North

Carolina, in a blueberiy patch near Lake
Michigan, from oak dunes in Indiana, by
beating underbrush in Arkansas, from a
roadcut in Missouri, in salt marshes, coastal
oak woods and by sweeping a meadow in
California. Judging by these notes, C. tur-
binata prefers more open areas than C.
conica, but according to Berry (1970)
Cyclosa turbinata has no clear habitat pref-
erences in North Carolina. H. K. Wallace's
field notes report specimens from a slope
near a sti'eam, the web attached to a stump,
from a stream bottom, from a slope near a
stream in Giles County, Virginia and in an
old field on a stream bank and in a sterile
area with fetterbush (Leucothoe) in Flor-
ida. I have collected specimens in central
Florida in dry grassy "prairie." Males are
matme from July to August in New York,
Pennsylvania and Virginia, from June to
September in the Southeast, from March to
August in Florida, to October in Texas, and
from March to September in California.
Females have been collected from May to
September in northern part of the range, in
all seasons except December to February in
Florida.

Distribution. Connecticut, cenb-al New
York, southern Michigan to Washington,
south to Central America, West Indies, and
also Bermuda, Cocos Island and Galapagos
Islands (Map 2).

Cyclosa caroli (Hentz)

Plate 3, Figures 51-63, Map 2

Epeira caroli Hentz, 1850, J. Boston Soc. Natur.
Hist, 6: 24, pi. 3, fig. 15, 2. Female type from
Alabama, destroyed. Keyserling, 1863, Sitzungs-
ber. Naturges. Isis Dresden, p. 137, pi. 6, figs.
17-19, 9.

Cyclosa laceria O.P.-Cambridge, 1889, Biologia
Centrali-Americana, Araneidea, 1: 50, pi. 7, fig.
14, S (as Epeira Jaccrta). Male lectotype here
designated from Guatemala or Panama in the
British Museum, Natural History, examined.
Keyserling, 1893, Spinnen Amerikas, 4: 275, pi.
14,' fig. 204, S . F.P.-Cambridge, 1904, Biologia
Centrali-Americana, Araneidae, 2: 494, pi. 47,
fig. 3, $ . NEW SYNONYMY.

Cijclosa caroli, — McCook, 1893, American Spiders,
3: 277, pi. 17, figs. 7, 8, 9, $. Keyserling, 1893,
Spinnen Amerikas, 4: 272, pi. 14, fig. 202, 9.

Cyclosa, Metazygia and Eustala • Levi

Figures 38-50 Cyclosa turbinata (Walckenaer): 38. Male from side. 39. Female from side. 40-43. Epigynum:
40 41 Ventral 42 43. Posterior. 40, 42. Cleared. 44-47. Male left palpus: 44. Apical. 45. Mesa!. 46.
Ventral. 47. Median' apophysis, ventral. 48. Female, dorsal. 49. Female abdomen, ventral. 50. Male, dorsal.

Scale lines. 0.1 mm, except Figs. 38, 39, 48-50, 1.0 mm.

$. F.P.-Cambridge, 1904, Biologia Centrali-

Americana, Araneidea, 2: 494, pi. 47, fig. 4, 9_.

Comstock, 1940, Spider Book, rev. ed., p. 467.

Roewer, 1942, Katalog der Araneae, 1: 761.

Bonnet, 1956, Bibliographia Araneorum, 2: 1310.
Cyclosa conigcra F.P.-Cambridge, 1904, Biologia

Centrali-Americana, 2: 494, pi. 47, fig. 5, 9.

Ten female syntypes from Omilteme, Mexico in

the British Museum, Natural History, examined.

NEW SYNONYMY.
Cyclosa elongate Franganillo, 1930, Mem. Inst.

Nac. Invest. Cient., 1: 68. Type specimens from
Sierra Maestra and Montanas de Trinidad in
Cuban Academy of Sciences, lost.
Parazygia accentonotata di Caporiacco, 1955, Acta
biol. Venezuelica, 1: 345, fig. 30, S. Male holo-
type from Rancho Grande, Aragua, \'enezuela
in the collections of Universidad Central, Cara-
cas, Venezuela, examined. NEW SYNONYMY.

Measurements. Female from Florida:
Total length 6.0 mm. Carapace 1.7 mm

84 Bulletin Mitseiun of Comparative Zoology, Vol. 148, No. 3

long, 1.1 wide. First femur, 1.4 mm; patella
and tibia, 1.7; metatarsus, 0.9; tarsus, 0.4.
Second patella and tibia, 1.4 mm; third, 0.9;
Fourth, 1.4.

Male from Florida: Total length 2.7 mm.
Carapace 1.4 mm long, 0.9 wide. First
femm-, 1.3 mm; patella and tibia, 1.2; meta-
tarsus, 0.7; tarsus, 0.4. Second patella and
tibia, 1.1 mm; third, 0.6; fourth, 1.1.

Variation. Females vaiy in total length
from 3.7 to 6.8 mm, carapace 1.3 to 1.9 long,
0.8 to 1.2 wide. Males vary in total length
from 3.0 to 3.4 mm, carapace 1.5 to 1.7 long,
1.1 to 1.2 wide. The largest individuals
came from Mississippi and Panama, the
smallest from Florida and Trinidad. The
tail of the female and especially of the male
xaries in length.

Diagnosis. Females of C. caroli found
north of Mexico can be separated from
other species by the shape of the abdomen
(Figs. 52, 61). The epigynal scape of C.
caroli is almost always oval to round (Fig.
54) and is lightest in the center, unlike the
scape of Central and South American spe-
cies with a similar abdomen. Male individ-
uals almost always have a small abdominal
tail (Fig. 51), lacking in C. turbinata males.
Males differ from related species also in the
shape of the short palpal median apophysis,
which has a distal hook and a convexly
curved distal keel below the hook (Figs. 59,
60). The middle spine present in C. ttir-
1)inata and C. tcalckenaeri median apophy-
sis is absent.

Natural History. Field notes of H. K.
Wallace report it from dense palmettos in
palmetto, in live-oak-hammock and in a ra-
\ine, both in Alachua Co., Florida. I have
collected specimens in Baygall woods and
mixed cypress forest in central Florida.
Comstock (1940) observed the species in a
"jungle near Miami, Fla. The orb of the
adult is six inches in diameter. The female
fastens her egg-sacs in a series which extend
across the web from the hub to the upper
margin like a stabilimentimi, and looks like
a dead twig caught in the web. This band

of egg-sacs and the spider are of the same
gray colour. When disturbed the spider
rushes to the band and appears as if it were
part of it. And here he will cling motion-
less even when the band is removed from
the web. ... I also observed smaller indi-
viduals shake their webs; these clung to the
stabilimentiuu, projecting the body at right
angles to it and in this position shook the
web violently." (Plate 3.)

Ruth Buskirk, in a note with the collec-
tions, says she found the "species very com-
mon in woods and woods edge in Costa
Rica. The orb has 25 radii, 22 spiral turns
... a radius of 8-12 cm, always vertical
orientation, debris and insect [remains]
wrapped with silk into long straight lines,
. . . 2 's often with egg cases in upper line."

Adult males have been collected in Feb-
ruary, July, September and December in
Florida, in June and July in Central Amer-
ica. Females are mature in all seasons.

Distribution. Georgia, Florida, Gulf
states, Mexico, Central America, West In-
dies, to southern Colombia, Venezuela and
Guyana (Map 2).

Cyclosa walckenaeri (O.P.-Cambridge)
Plate 4, Figures 64-77, Map 2

Epeira bifurcata, — Keyserling, 1863, Sitzungsber.
Natiirf. Gesell. Isis, Dresden, p. 142, pi. 6, figs.
22-23, 2 . Specimens from Bogota, Colombia.
Not Epeira bifurcata Walckenaer, 1841.

Turckheimia ivalckenaerii O.P.-Cambridge, 1889,
Biologia Centrali-Americana, Araneidea, 1: 47,
pi. 8, fig. 6, 2 . Three female syntypes from
Volcan de Fuego, Guatemala in the British Mu-
seum, Natiual History, examined.

Epeira walckenaerii Keyserling, 1892, Spinnen
Amerikas, 4: 98, pi. 5, fig. 73, 9, $. Types
from Bogota, Colombia, Guatemala, Taquara do
Mundo novo and Rio Grande do Sul, Brazil in
the British Museum, Natural History.

Cyclosa walckenaeri, — McCook, 1893, American
Spiders, 3: 226, pi. 17, fig. 1, $, £. F.P.-Cam-
bridge, 1904, Biologia Centrali-Americana, Ara-
neidea, 2: 495, pi. 47, fig. 9, $. Petrunkevitch,
1930, Trans. Connecticut Acad. Sci., 30: 315,
figs. 188, 189, 9 .

Cyclosa trifida F. P. -Cambridge, 1904, Biologia
Centrali-Americana, Araneidea, 2: 495, pi. 47,
fig. 7, 2 . Three female syntypes, slightly dam-

Cyclosa, Metazygia and Eustala • Levi

Figures 51-63. Cyclosa caroli (Hentz): 51. Male from side. 52. Female from side. 53-56. Epigynum; 53, 54.
Ventral. 55, 56. Posterior. 53, 55. Cleared. 57-60. Male left palpus: 57. Apical. 58. Mesal. 59. Ventral.
60. Median apophysis. 61. Female, dorsal. 62. Female abdomen, ventral. 63. Male, dorsal.

Scale lines. 0.1 mm except Figs. 51, 52, 61-63, 1.0 mm.

aged from Cohabon, Guatemala, in the British
Museum, Natural History, examined. NEW
SYNONYMY.
? Cyclosa ciiadritubcwsa Franganillo, 1936. Ardc-
nidos de Cuba, p. 84. Juvenile liolotype from
Cuba in the Cuban Academy of Science, in poor
condition, examined. It appears to lack lateral
posterior tubercles.

Note. Specimens of tliis species and
several similar South American species in
both the American Museum and the Mu-
seum of Comparative Zoology had been la-

beled Cyclosa oculata. Cyclosa oculata
(Walckenaer) (Figs. 21-23) is a Em-opean
species not found in the Americas. The
type specimens of the name came from
Paris. This error dates from Simon (1900),
who listed C. oculata as occurring in Hawaii,
the United States, Antilles and X^enezuela
and indicated that Epeira tcalckenaeri Key-
serling is probably a synonym. Simon did
not examine genitalia carefully and the
shape of the abdomen of the two species

Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

is similar. E. B. Bryant (1940), skeptical
of the synonymy, borrowed specimens of
C. oculata from Paris and got specimens
determined by Simon which were the same
species as C. walckenaeri. Not surprisingly,
they came from America: Hispaniola.

The three syntypes of C. trifida have the
characteristic epigynum but the abdomen
is flattened, apparently damaged when col-
lected. They have the four posterior tuber-
cles, but not the two anterior ones.

Measurements. Female from Texas: To-
tal length 6.3 mm. Carapace 2.2 mm long,
1.5 wide. First femur, 2.0 mm; patella and
tibia, 2.2; metatarsus, 1.1; tarsus, 0.6. Sec-
ond patella and tibia, 1.9 mm; third, 1.0;
fourth, 1.7.

Male from Texas: Total length 3.8 mm.
Carapace 1.7 mm long, 1.4 wide. First
femur, 1.7 mm; patella and tibia, 1.7; meta-
tarsus, 1.0; tarsus, 0.6. Second patella and
tibia, 1.2 mm; third, 0.7; fourth, 1.4.

Variation. Total length of females 3.8 to
6.8 mm, carapace 1.3 to 2.0 long, 1.0 to 1.2
wide. Total length of males 2.1 to 3.8 mm,
carapace 1.1 to 1.7 long, 0.8 to 1.3 wide.
The smallest females came from Florida, the
largest individuals from Guatemala. Some
specimens have the abdomen longer. Rarely,
the sides of the scape of the epigynum are
curved out and the scape slightly oval. One
female from Oriente Province, Cuba had an
epigynum like that of C. walckenaeri, but
the abdomen was like that of C. caroli, with
only faint indications of humps.

Diafinosis. The four humps on the poste-
rior tip of the abdomen and two dorsal
humps anterior of the middle separate the
species from other Cijclosa in North Amer-
ica. Unlike C. caroli and C. turhinata the
sides of the epigynum scape are usually
parallel, making it a narrow rod (Fig. 67).
Males can usually be readily separated by
the indications of the four posterior abdo-
men humps (Figs. 64, 76). The median
apophysis of the palpus is very long, but
unlike that of C. turhinata, the middle spine
is minute and the distal hook continues into

a keel proximally (Figs. 72, 73). That of C.
caroli lacks the middle tooth entirely and is
short.

Natural History. Specimens of C. icalck-
enaeri have been found on large aloe and in
open shi-ubs at edge of woods in Jamaica, in
dry sluubs in the Virgin Islands, in a garden
in Cuba, on shrubby edge of woods along
coast of Florida Keys, on mangroves in Baja
California and in a pine-oak forest in Chia-
pas. The eggs are hung in the web. Webs of
juveniles observed in Florida had a narrow
stabilimentimi of debris (Plate 4) and the
only one containing egg-sacs had been
destroyed and left unfinished.

Males have been collected in May, Au-
gust, September and October in the south-
ern states and northern Mexico and females
in all seasons.

Distribution. Southern Florida, southern
Texas, central California coast to Panama
and West Indies (Map 2).

Cyclosa bifurca (McCook)

Plate 5, Figures 78-89, Map 2

Cyrtophora bifurca McCook, 1887, Pioc. Acad.
Natur. Sci. Pliiladelpliia, 3: 342. Female, male
syntypes from Fairyland, Merrit's Island on the
Indian River, Florida in the Philadelpliia Acad-
emy of Sciences, lost.

Ctjclosa fissicauda O.P.-Cambridge, 1889, Biologia
Centrali-Americana, Araneidea, 1: 49, pi. 8, fig.
7, $ . Fifteen syntypes in two vials, from near
Dolores, Guatemala in the British Museum,
Natural History, examined. Ke>serling, 1893,
Spinnen Amerikas, 4: 274, pi. 14, fig. 203, 9.

Cyclosa bifurca, — McCook, 1893, American Spiders,
3: 227, pi. 17, figs. 9, 10, 9,6. F.P.-Cam-
bridge, 1904, Biologia Centrali-Americana, Ara-
neidea, 2: 495, pi. 47, fig. 8. Comstock, 1940,
Spider Book, p. 467, figs. 465, 466, ? , egg-sacs.
Roewer, 1942, Katalog der Araneae, 1: 759.
Bonnet, 1956, Bibliographia Araneorum, 2(2):
1309.

Description. Female from Florida: In
alcohol, carapace yellow-wliite, sternum
brown with a central longitudinal white
band and white patches near base of ante-
rior three coxae. Leo;s vellow-white with
some indistinct dark bands distally. Dor-
sum of abdomen white with indistinct

Cyclosa, Metazygia and Eustala • Levi

Figures 64-77. Cyclosa walckenaeri (O.P.-Cambridge): 64. Male from side. 65. Female from side. 66-69.
Epigynum: 66, 67. Ventral. 68, 69. Posterior. 66, 68. Cleared. 70-73. Male left palpus: 70. Apical. 71.
Mesal. 72. Ventral. 73. Median apophysis. 74. Female, dorsal. 75. Female abdomen, ventral. 76. Male,
dorsal. 77. Male coxae.

Scale lines. 0.1 mm except Figs. 64, 65, 74-77, 1.0 mm.

88 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

marks, sides with indistinct gray marks.
\^enter with a white square whose sides are
lateral to the spinnerets. The legs are thick
( Fig. 88 ) . Total lengtli 6.5 mm. Carapace
2.2 mm long, 1.7 wide. First femur, 2.5 mm;
patella and tibia, 2.7; metatarsus, 1.6; tarsus,
0.8. Second patella and tibia, 2.2 mm; third,
1.2; fourth, 2.1.

Male from Florida: Carapace and abdo-
men yellowish white with a median black
longitudinal line on carapace, some indis-
tinct black pigment spots on the abdomen.
Posterior median eyes 0.6 diameter of ante-
rior medians. Anterior laterals 0.5, posterior
laterals 0.6 diameters. Anterior median eyes
their diameter apart, 0.7 from laterals. Pos-
terior median eyes their diameter apart, 1.5
from laterals. Neither coxae nor legs modi-
fied. The abdomen is like that of female, but
the humps are barely visible. Total length
1.8 mm. Carapace 0.9 mm long, 0.7 wide.
First femur, 1.0 mm; patella and tibia, 1.1;
metatarsus, 0.9; tarsus, 0.4. Second patella
and tibia, 0.8 mm; third, 0.4; fomth, 0.6.
Another male measured 1.7 mm total length.

Note. The live spider and the egg-sacs
are green, the venter of tlie abdomen having
a bright red patch between epigynum and
spinnerets (Comstock, 1940). The color
washes out in alcohol. The egg-sac is an ir-
regular octagon, and as many as 10-14 egg-
sacs may be strung together (McCook,
1887). The male is minute. Only one male
was found in a collection of 207 specimens.
About another 130 specimens yielded only
one more male.

Variation. Total length of females 5.1 to
9.0 mm long, carapace 2.0 to 2.9 mm long,
1.5 to 2.3 mm wide. Some individuals have
more black pigment than others and have
the legs ringed.

Ditt'^nosis. North of Mexico no other
American species of Cijclosa has a forked
tail (Figs. 78, 88). Ctjclosa furcata O.P.-
Cambridge is similar in appearance but the
epigynum has a scape and the base differs
in shape.

Natural History. J. Boursot collecting in
El Salvador reported on notes in the vial:

"with contracted legs these spiders crouch
at one end of the stabilimentum composed
of rejected chewed food wliich they match
identically. Discovered only on tarred sur-
face of huge water tank." C. B. Worth
( 1940 ) reported on the shape and coloration
of the animals whose vertical webs, six
inches in diameter, he saw on the walls of a
Florida house, parallel to the walls: "The
egg-sacs are arranged in a row, occupying
the position of the hands of a clock at ex-
actly noon. The spider herself reposes at
the center of the web, that is immediately
below and toucliing the lowermost egg-sacs.
She invariably faces the ground, so that her
abdomen appears as an additional egg-sac
in the row above her. . . ." The "mass of ob-
jects in the web is that of a catkin. . . . This
appearance is heightened by the spider's
disposition of captured food. Such prey is
wrapped in silk and anchored below the
spider, forming an uneven row of objects as
a direct short continuation of the line of
egg-sacs. The average length of the 'cat-
kins,' i.e. egg-sacs, spider food-sacs ... is
from two-and-a-half to three inches, which
means that they occupy about half the
diameter of the web. The usual number of
egg-sacs ranges from five to nine with eight
on an average. But the most remarkable
feature of all is the resemblance of the egg-
sac to the abdomen of the female. The
latter is light green with dark green central
and lateral stripes and in these details the
egg-sacs agree precisely with theii- maternal
source. The spider's abdomen moreover
bears a series of tubercles and projections,
which again are reproduced faithfully in the
egg-sacs even inclucUng the terminal bifur-
cation. The egg-sacs are finally deposited
in the web in a shingled or overlapping
series, and the spider takes a position at the
center of tlie web so that her abdomen over-
laps the lowermost egg-sac in an exact con-
tinuation of the series above her. . . . The
spider's light green color and smooth integu-
ment give it a translucent appearance when
seen close at hand. Even tliis quality of

Cyclosa, Metazygia and Bust ALA • Levi

Figures 78-89. Cyclosa bifurca (McCook): 78. Female from side. 79-85. Epigynum: 79, 80, 84. Ventral. 81,
82, 85. Posterior. 83. Lateral. 79, 81. Cleared. 79-83. (Florida). 84,85. (Texas). 86,87. Male left palpus:
Se'. Mesal. 87. Ventral. 88. Female, dorsal. 89. Male, dorsal.

Scale lines. 0.1 mm except Figs. 78, 88, 89, 1.0 mm.

translucence is duplicated in the smooth-
woven texture of the egg-sacs." (Plate 5.)
The spider has been collected on a torn-
down building in a wooded area of Austi-a-
lian pines {Casuarina sp.) and cabbage
palms ( Sahal palmetto ) and on a saw pal-
metto leaf (Serenoa sp.) in Florida, from
the nest of a wood rat (Neotoma sp.), and
from a wasp nest. One record is from an
arid, subtropical area in San Luis Potosi.
Comstock ( 1940) found it in a "jungle near-
the shore" and also on the "ceiling of a

veranda by the hundred." Matiu-e females
have been collected in every month in Flor-
ida and Texas.

Distribution. Florida, southern Alabama,
soutliern Texas, Mexico to El Salvador,
Cuba and Hispaniola (Map 2).

Metazygia F.P.-Cambridge

Metazygia F.P.-Cambridge, 190.3, Biologia Cen-
tral!-Americana, Araneidea, 2: 501. Type spe-
cies by original designation M. icittfeldae
(McCook). The name is feminine.

Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Diagnosis. The abdomen is spherical
(Fig. 108) to round and dorsoventrally
flattened (Figs. 98, 109) as in Niictenea and
Zijgiella, but differs from those two genera
by having no pigment ventrally between
genital groove and spinnerets (Figs. 99,
117). The carapace differs from that of
Niictenea by lacking fine setae (Figs. 96,
108), and the epigynum differs by lacking
a scape. In place of the scape is a laterally
flattened knob (Figs. 90-92), which can
expand and project anteriorly in M. zilloides
(Figs. 104-106) so as to resemble the
epigynum of Eustala species. There is no
such knob in M. carolinalis (Fig. 112).

Males differ from Nuctenea in having
only one macroseta on the palpal patella,
as in Zijgiella, and differ from Zijgiella in
the very different structiu'e of the palpus.
Metazijgia, unlike Zijgiella, has a hook-
shaped paracymbium (p in Fig. 103), a
transparent subterminal apophysis (sa),
and a knob-shaped median apophysis ( m ) ,
ventrally attached (Figs. 101-103). Meta-
zijgia resembles Zijgiella in having the tegu-
lum (t) of the palpus modified; however,
the modification is apical (Figs. 102, 111).
The palpus is similar to that of Eustala but
the median apophysis (m) is always knob-
shaped (Figs. 101-103, 110, 111), not cone-
shaped as in Eustala.

Description. Carapace smooth with few
hairs, often darker anteriorly than poste-
riorly (Fig. 96), or with a median longi-
tudinal pigment line (Fig. 108), wdth little
or no thoracic depression.

Eye sizes subequal (M. carolinalis) or
anterior median eyes slightly larger than
others (M. wittfeldae, M. zilloides). Later-
als some distance from medians (Fig. 97)
except in the small M. zilloides in which the
eyes of the anterior row are equally spaced.
Height of clypeus slightly less than diam-
eter of anterior median eyes (Fig. 97).
Chelicerae very strong, bulging proximally
(Fig. 97), narrower distally, especially in
M. carolinalis. Legs tliick and strong (Figs.

96, 108), not banded, with many macrosetae
and setae. First leg longest, legs 1,2,4,3.
Abdomen oval to round, more or less dorso-
ventrally flattened (Figs. 96, 98, 108, 109,
116). In M. carolinalis the abdomen has
dorsal sclerotized discs (Fig. 116). No
black pigment on venter (Figs. 99, 117).

Males slightly smaller (Fig. 100) than
females, with similar coloration and eyes.
The chelicerae and fangs of some ti'opical
species are modified, perhaps for copula-
tion. Legs differ from those of females by
being slightly longer and having more
macrosetae (Fig. 100), especially on the
second tibia. The distal margin of first coxa
has a hook that fits into a groove on the
second femur.

Genitalia. The base of the epigynum has
a ventral, laterallv compressed knob in
place of the scape '(Figs. 90-92, 104-106);
the knob is absent in M . carolinalis; in M.
zilloides it projects anteriorly if expanded,
resembling that of Eustala.

The male palpus, similar to that of Eus-
tala, differs in several ways. The tenninal
apophysis is a prong (a in Figs. 103, 110,
111), the subterminal apophysis often a
transparent bubble (sa in Figs. 101-103,
110). The embolus (e), hidden in the tem-
perate species, may have a piece that
breaks off during mating and (in M. zil-
loides) remains in the epigynum. (But
this is not certain, as the two common spe-
cies north of Mexico have the embolus hid-
den behind the conductor and subterminal
apophysis.) The conductor is a complex
sclerite and the median apophysis (m) a
simple knob (Figs. 102, 103, 110, 111), not
a cone hanging down as in Eustala. The
Metazijgia palpus has a large sclerite me-
sally wliich may be the stipes (Figs. 101,
103, with texture in 110); it differs in shape
in related tropical species.

Natural History. Unlike the related Eus-
tala, Metazijgia makes a reti'eat near the orb
web. Metazijgia wittfeldae is often found
on bridges and buildings; Metazijgia witt-

Cyclosa, Metazygia and Eustala • Levi

Metazygia zilloides

Map 3. Distribution of Metazygia carolinalis (Archer), M. wittfeldae (McCool<) and M. zilloides (Banks).

feldae becomes active after dark, tearing
down remnants of the old web and making
new radii, scaffolding and viscid threads.
The old web is usuallv left nntil a new one
is built, which may not be every night.
Threads coated with cornstarch ( dusted by
photographers the previous night) are
hauled in, two sections at a time, balled up,
and thrown horizontally away from the
web, with some force, at the rate of a ball
every minute or two. Silk not dusted is
probably eaten. During the day the spider
remains in a crevice; at night it hangs in the
center of the web. The light from a flash-
light may cause the spider to move away.
The webs observed at the Archbold Bio-
logical Station, Lake Placid, Florida were
loose with few frame threads and 10 to 18

radii. The number of viscid threads in
several webs was 18, 16, 22, 25, 15 below
the hub and 11, 3, 3, 17, 10 above the hub.
The webs had solid hubs (Plate 6) and
horizontal diameters ranging from 10 to 27
cm. \\xbs, as many as five next to each
other in a suitable comer, were vertical be-
tween railings of a ramp 35 cm above the
ground; at 2.7 m above the level of the
ramp, imder the ceiling, the webs were al-
most horizontal. The spiders avoided the
area near a light fixtine, but used areas
some distance away, where they hai"vested
insects attracted to the light.

Species. There are three species north of
Mexico; most other species are tropical
American (Map 3); none is known outside
of America.

Bulletin Miiscuiii of Comparative Zoology, Vol. 148, No. 3

Key to female Metazygia
1. Epigynum without \entral median knob
(Fig. 112); dorsum of abdomen with 4
pairs of sclerotized discs (Fig. 116); North
CaroHna carolinalis

— Epig\num with a ventral median, laterally

compressed knob (Figs. 90-92, 104-106);
abdomen without sclerotized discs; Vir-
ginia south to Texas 2

2(1) Median knob very narrow, areas to side
and anterior to it soft and expandable
( Fig. 104 ) ; openings of epigynum on ven-
tral face on each side (Fig. 104); dorsum
of abdomen with a pair of anterior black
marks (Fig. 108) zilloides

— Median knob wide; areas to side and anterior

to it not expandable (Fig. 90); openings
of epigynum posterolateral of base (Figs.
91, 92); dorsum of abdomen with a series
of dark brackets, farthest apart anteriorly,

and a median dark line (Fig. 96)

wittfeldae

Key to male Metazygia

(M. carolinalis male unknown)

1. Terminal apophysis prong of palpus pointed

(Figs. 101-103) wittfeldae

— Terminal apophysis prong of palpus with

blunt tip, wider at tip than proximally
(Figs. 110, 111) zilloides

Metazygia wittfeldae (McCook)
Plate 6, Figures 90-103, Map 3

Epeira wittfeldae McCook, 1893, American Spiders,
3: 168, pi. 7, figs. 6, 7. Three female, two male
and one male juvenile syntypes from Florida in
the Academy of Natural Sciences, Philadelphia,
examined.

Metazygia ivittfeldae, — F.P.-Cambridge, 1904, Bio-
logia Centrali- Americana, Araneidea, 2: 501, pi.
47, figs. 22, 23, 9, S. Roewer, 1942, Katalog
der Araneae, 1 : 868. Bonnet, 1957, Bibliographia
Araneorum, 2(3): 2820.

Description. Female from Florida: Cara-
pace with head region much darker brown
than thorax ( Fig. 96 ) . Sternum, legs orange.
Dorsum of abdomen light brown with pairs
of dark marks approaching each other
posteriorly (Fig. 96). Total length 8.0 mm.
Carapace 4.2 mm long, 3.0 wide. First
femur, 3.6 mm; patella and tibia, 4.0; meta-
tarsus, 2.7; tarsus, 1.2. Second patella and
tibia, 3.7 mm; third, 2.3; fourth, 2.9.

Male: Total length 5.8 mm. Carapace

3.5 mm long, 2.4 wide. First femur, 3.6 mm;
patella and tibia, 4.4; metatarsus, 3.4; tar-
sus, 1.4. Second patella and tibia, 4.0 mm;
third, 2.2; fourth, 2.7.

Variation. Females varied in total length
from 6.0 to 10.2 mm; carapace 2.9 to 4.2
long, 2.5 to 3.4 wide. Males varied, total
length 5.0 to 7.0 mm; carapace 3.0 to 4.0
long, 2.2 to 3.1 wide.

Diagnosis. Females of M. wittfeldae dif-
fer from a similar West Indian species and
from M. duhia (Keyserling) in Central and
South America by the epigynum, which, in
posterior view, has overhanging lateral
bulges of the median area (Figs. 91, 92).
Males differ by having tlie embolus hidden
by the large subterminal apophysis (Figs.
101-103 ) , a tooth at the base of the conduc-
tor (c in Figs. 102, 103) and a pocket at the
distal edge of the tegulum (t in Figs. 102-
103).

Natural History. This species is com-
monly found under the eaves of buildings
from \^irginia to Florida, and also on houses,
and on and under bridges. In Florida, it has
been found in cypress swamp, in tall grass,
in citrus tree foliage, in vegetation border-
ing a canal, on canal banks with heavy cut
grass and ragweed, and on slash pine {Pinus
elliottii). Many specimens came from wasp
nests. The web (Plate 6) is described above
in the introduction to the genus Metazygia.

Distribution. From Norfolk, Virginia (nu-
merous collections from buildings around
Stumpy Lake) to Florida, Gulf states to
Texas to Centi-al America (Map 3).

Metazygia zilloides (Banks), new
combination

Figures 104-111, Map 3

Epeira zilloides Banks, 1898, Proc. California Acad.
Sci., 3 ser., 1: 255, plate 15, fig. 2, $, S. Three
female, one male, one juvenile syntypes from
Tepic, Mexico in die Museimi of Comparative
Zoology, examined.

Arauca dilatata F.P.-Cambridge, 1904, Biologia
Centrali- Americana, Araneidea, 2: 513, pi. 49,
fig. 9, $ . Male lectotjpe here designated from
[no locality] Guatemala in the British Museum,
Natural History, examined. There are three

Cyclosa, Metazygia and Evstala • Levi 93

Figures 90-103. Metazygia wittfeldae (McCook): 90-94. Epigynum: 93-95. Cleared. 90, 93. Ventral. 91, 94.
Posterior. 92, 95. Lateral. 96. Female, dorsal. 97. Female, eye region and chelicerae. 98. Female from
side. 99. Female abdomen, ventral. 100. Male from side. 101-103. Left male palpus: 101. Mesal. 102.
Ventral. 103. Mesoventral, expanded.

Abbreviations, a, terminal apophysis; c, conductor; dh, distal hematodocha; e, embolus; m, median apophy-
sis; p, paracymbium; r, radix; sa, subterminal apophysis; t, tegulum.

Scale lines. 0.1 mm, except Figs. 96-100, 1.0 mm.

94 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

paralectotvpes, of which one is Metazygia in-

certa. NEW SYNONYMY.
Metazygia keyserlingi Banks, 1929, Bull. Mus.

Comp. Zool., 69: 94, fig. 63. Female holotype

from Barro Colorado Island, Canal Zone, in the

Museum of Comparative Zoology, examined.

NEW SYNONYMY.
Metazygia alhonigra, — Biyant, 1940. Bull. Mus.

Comp. Zool., 86: 339, figs. 107-109, 111, $. $,

erroneous determination, not Lamia alhonigra

Franganillo.
Aranetis pallidulus, — Kraus, 1955, Abhandl. Senck-

enbergischen Naturf. Gesell., 493: 24, fig. 66,

5 . Erroneous detennination.

Note. American Museum specimens had
been labeled Metazygia incerta, Museum of
Comparative Zoology West Indian speci-
mens as Metazygia alhonigra (Franganillo)
and Florida and Texas specimens as Epeira
pallidula (Keyserling) by Biyant, and as M.
keyserlingi by Chickering. The name Meta-
zygia incerta belongs to a different species.
The name Larinia all)onigra is a synonym of
L. directa and the specimens were incor-
rectly determined by Bryant. M. keyserlingi
is a synonym of M. zilloides.

Description. Female from Florida: Cara-
pace light yellowish brown with a narrow,
median, longitudinal dark band on cara-
pace. Sternum, legs, light brownish. Dorsum
of abdomen white with anterior pair of dark
patches and four pairs of dark spots (Fig.
108). Total length 6.1 mm. Carapace 2.3
mm long, 1.8 wide. First femur, 2.2 mm;
patella and tibia, 2.7; metatarsus, 1.8; tar-
sus, 0.8. Second patella and tibia, 2.2 mm;
third, 1.3; fourth, 2.0.

Male from Florida: Total length 4.0 mm.
Carapace 2.2 mm long, 1.7 wide. First
femur, 2.7 mm; patella and tibia, 3.5; meta-
tarsus, 2.7; tarsus, 1.0. Second patella and
tibia, 2.8 mm; third, 1.4; fourth, 2.0.

Variation. Some specimens have the pos-
terior of the abdomen dark and there are
wliite rings around the black spots. Total
length of females 3.6 to 7.4 mm, carapace
1.8 to 3.2 long, 1.4 to 2.4 wide. Total length
of males 3.4 to 4.8 mm, carapace 1.7 to 2.6
long, 1.3 to 2.0 wide. Males from Cuba

have the distal edge of die tegulum smooth,
with no teeth.

Diagnosis. Most specimens have the an-
terior black patches on the abdomen and a
series of dark spots (Fig. 108). Females
differ from both M. wittfeldae and M. in-
certa (O.P.-Cambridge) by having antero-
ventrally directed openings on each side of
the epigynal base (Fig. 104). Males differ
from M. wittfeldae by the blunt terminal
apophysis (Figs. 110, 111), and from M.
wittfeldae and M. incerta by die shape of
the (textiued) stipes (Fig. 110), the shape
of the conductor ( under terminal apophysis.
Fig. Ill), and the toothed edge on the distal
surface of the tegulum (Fig. 111).

Natural History. The species has been
collected by sweeping flowers in Texas,
in Florida in palmetto-poisonwood flats,
among roadside weeds along a canal, in
shrubs and vegetation, and on Casiiarina
(Australian pine). Males have been col-
lected in Florida in June.

Distribution. Southern Florida, central
and southern Texas to Colombia; Cuba,
Jamaica and Trinidad (Map 3).

Metazygia carol! nails (Archer), new
combination

Figures 112-117, Map 3

Epeira carolinalis Archer, 1951, Amer. Mus. Novi-
tates, no. 1487: 40, fig. 57, 9. Female holotype
from White Lake, Bladen County, North Caro-
lina, in the American Museum of Natural His-
tory, examined.

Description. Female: Carapace dark
brown on sides, brown above. Legs brown.
Sternum light brown. Dorsum of abdomen
with sclerotized discs brown, wliite pigment
spots, and two dark lines, one on each side,
approaching each other anteriorly and pos-
teriorly (Fig. 116). Venter with a pair of
indistinct white brackets, no black pigment
(Fig. 117). The carapace is flat and very
low (Fig. 116). Abdomen oval, dorsoven-
trally flattened (Fig. 116). Total length
11.0 mm. Carapace 4.5 mm long, 3.7 wide.
First femur, 3.7 mm; patella and tibia, 5.5;

Cyclosa, Metazygia and Eustala • Levi 95

Figures 104-111. Metazygia zilloides (Banks): 104-107. Epigynum: 104. Ventral. 105. Posterior. 106. Lat-
eral. 107. Posterior, cleared. 108. Female, dorsal. 109. Female, lateral. 110, 111. Left male palpus: 110.
Mesal. 111. Ventral.

Figures 112-117. Metazygia carolinalis (Archer): 112-115. Epigynum: 112. Ventral. US. Posterior. 114.
Lateral. 115. Dorsal, cleared. 116. Female, dorsal. 117. Female abdomen, ventral.

Scale lines. 0.1 mm except Figs. 108, 109, 116, 117, 1.0 mm.

Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

metatarsus, 3.7; tarsus, 1.4. Second patella
and tibia, 4.8 mm; third, 2.7; fourth, 4.3.

Diagnosis. Unlike other Metazygia spe-
cies, M. coroUnaJis lacks a ventral knob
(Fig. 112) on the epigynum.

Note. The placement of this species in
Metazygia is doubtful. Archer thought M.
caroUnalis close to Niictenea cornuta and
placed it with cornuta in Epeira. He may
have been right. But the following facts
speak against this placement. Niictenea is
mainly a Palearctic genus with a few species
in North America having a holarctic dis-
tribution. One of the main characters of
Nuctenea females is the black venter with
the comma-shaped wliite marks on each
side. This is not present in M. caroUnalis.
The placement of the species will remain
uncertain until the male is found.

Natural History. The flattened shape of
the spiders, especially the low carapace,
suggests that the spider has its retreat in
crevices, probably under bark.

Records. North Carolina: Bladen Co.,
$ paratyi3es, Sept. 1929 (J. C. Beakley);
Craven Co.: New Bern, May 1900, 2 ?, 1
juv. ( J. H. Emerton) (Map 3).

Eustala Simon

Eiistah Simon, 1895, Histoire Naturelle des Araig-
nees, 1: 795. Type species Epeira anastera
Walckenaer by original designation; The name
is feminine.

Diagnosis. Eustala differs from other
Araneidae, especially from Araneus, by the
epigynum, which has its scape projecting
anteriorly (Figs. 118, 138, 140) instead of
posteriorly as in all other genera, and by the
male palpus, which has only one patellar
macroseta, and has the median apophysis,
a white cone-shaped structure, hanging
down the venter of the palpus (Figs. 126,
147, m in Fig. 232).

The carapace has a deep longitudinal
cleft in the thoracic region (Figs. 163, 183,
197). The abdomen is usually triangular,
pointed above the spinnerets (Figs. 142-
144, 209-210). Like Larinia and Metepeira,

but unlike many other Araneidae genera,
Eustala has a central, ventral white patch on
the abdomen (Figs. 155, 173, 185, 211).
The white patch is absent in those tropical
Eustala that have the abdomen elongate,
like that of Larinia. Juvenile Eriophora,
which look like Eustala, lack the white
patch and have a dark trapezoid on the ven-
ter.

The related Metazygia has the scape of
the epigynum projecting ventrally (Figs.
90, 91) and the median apophysis is a soft
knob (Figs. 101-103, 110, 111). The cara-
pace is smooth (Figs. 96, 108), and the ab-
domen is oval, slightly flattened dorsoven-
trallv, with indistinct ventral markings
( Figs. 96, 98, 99, 108, 109).

Description. The carapace is shaped as
in Araneus, but with a deep longitudinal
thoracic cleft (Figs. 163, 183, 197). The
carapace is covered with setae and the
thoracic area is high in some species (Figs.
133, 154, 172). The posterior median eyes
are slightly smaller than the anterior me-
dians, sometimes equal, rarely slightly
larger. The laterals are always smaller than
the medians. Anterior medians are their
diameter apart, or 1.5 diameters at most;
the posterior medians are separated by
about the same distance. The laterals (ex-
cept in the smallest species) are two to
several diameters from medians. The clyp-
eus height equals the diameter of the
anterior median eyes (Fig. 225) except in
E. clavispina where it is about one and one-
half the diameter of the anterior median
eyes as a result of the projection of the eye
area. There often is a dark transverse band
between anterior median and anterior lat-
eral eyes (Figs. 163, 210, 22.5). The legs
are more or less banded. The abdomen is
generally triangular with a posterior hump
^Figs. 209, 210), but this may be absent
(Figs. 122, 123, 257, 258) or there may be
several humps (Figs. 163, 164, 196, 197, 223,
224). Most species are variable in colora-
tion with dark and light individuals, but
most have a folium pattern on the dorsum,

Cyclosa, Metazygia and Eustala • Levi

exceptions being some specimens of E.
anastcia that are contrastingly colored with
black patches on white in alcohol (Figs.
219, 222). In most Eustala species, unlike
most species of Araneus, the venter has a
more or less distinct median ventral white
patch (Figs. 185, 198). In a few species
this white patch is as distinct and conti^ast-
ing (Fig. 173) as in Metepeira. Living
specimens of E. anastera from central Flor-
ida ha\'e a greenish abdomen, but the green
washes out of alcohol-presei'ved specimens.

Males are smaller than females, slightly
darker in color, their abdoininal humps are
less distinct tlian in females (Figs. 199,
212). The distal margin of the first coxa
has a hook (Fig. 201) which fits into a
groove on the second femur. Except for
being longer and having stronger macro-
setae, especially on the second tibia, the
legs of Eusfola are not modified. Some spe-
cies have a ventral row of macrosetae on
one or more femora (Figs. 125, 156, 189,
214). This is a species characteristic and
has been illustrated. The males are exceed-
ingly difficult to match with females: spe-
cies with the (seemingly) most specialized
palpi do not necessarily have the most
specialized epigvna (e.g. E. californiensis,
Figs. 138-148).'

Genitalia. The epigynum has an unusual,
anteriorly projecting scape, annulate in most
species but smooth in E. devia (Fig. 118)
and E. cazieri (Fig. 128). The three plates
in posterior view of the epigynum are of
diagnostic importance; the median and two
laterals, varying in shape. The seminal
receptacles are usually spherical; between
the openings is another smaller spherical
structure which appears to contain a wind-
ing duct (Figs. 208, 256).

The palpal patella has one macroseta
(Figs. 217, 252). The bulb, which is similar
to that of Metazygia, has a huge conductor
(c), variously shaped in different species,
and a \\'hitc, soft, conical median apophysis
(m), which hangs down on the venter of
the bulb in all Eustala species (Fig. 232).

The embolus (e) is a hook, similar in all
species, and has a large sclerotized base,
the stipes. The tenninal apophysis is a
sclerotized prong (a), slightly different in
different species, resting on a bubble-like,
transparent, spherical subterminal apophy-
sis (a in Fig. 232). In some species the ter-
minal apophvsis is different in shape ( Figs.
126, 136, 147, 157). The mesal side of the
palpus faces ventrally, the ventral side
laterally in resting position (Fig. 231).

Natural History. Considering the com-
mon occurrence of many Eustala species,
sui"prisingly little was known about them.
Eustala apparently is noctvunal and removes
its web at daytime. During the day it rests
on a dead branch; there is no retreat. Eus-
tala species are commonly collected by
sweeping and are found also as prey in
mud-dauber wasp nests.

Eustala anastera obsei^ved at the Arch-
bold Biological Station, Lake Placid, Flor-
ida made webs every evening after dark.
The webs usually had disappeared by the
morning, but once in awhile the webs are
not taken down. The webs of juveniles had
17 to 25 radii, that of an adult, 18 and 21.
The webs of tliese juveniles had 28, 37, 41,
31 and 15 viscid threads below the hub and
above the hub had 36, 38, 39, 28, 32. The
web of an adult had 30, 33 below, 31, 28
above. The horizontal diameter of juveniles'
webs ranged from 12 to 25 cm; of adults'
webs 19 and 30 cm. There were few frame
threads, the hub was solid (Plate 7). The
webs were built in dead branches, usually
away from leaves and within a wire fence,
having veitical wires 15.5 cm apart. Eustala
has no retreat; when not in the center of the
web, it sits appressed to branches. Most
webs are vertical but a horizontal web was
seen. The lowest webs are 3 to 4 feet above
tlie ground; the maximum height is not
known.

Eustala anastera in central Florida feeds
on a wide \'ariety of medium-sized prey,
and when resting in the web usually keeps
its legs slightly spread like Eviophora ravilla.

98 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Map 4. Distribution of Eustala devia (Gertsch and Mulaik), E. cazleri n. sp., E. californiensis (Keyserling), E.
bifida F.P.-Cambridge, E. brevispina Gertsch and Davis, E. clavispina (O.P.-Cambridge), E. cameronensis
Gertsch and Davis and E. eleuthera n. sp.

Cyclosa, Metazygia and Eustala ' Levi 99

Eustala rosae

Eustala con

/ i

'y<S:^M 1

^^-<-'-< ^^

Eustala anastera

/:-^

7TC — ; — \ —

• u^'^ • ^

•..V- :-•....(

Eustala cepina

I ^

..|3

r-7.-\ .

-V

^ 1 •/ v-

'^^-j

Eustala emertoni

Map 5. Distribution of Eustala rosae Chamberlin and Ivie, E. anastera (Walckenaer), E. cepina (Walckenaer),
E. emertoni (Banks), E. conchlea (McCook).

but unlike many other genera (M. Stowe,
personal communication ) .

Species and Distribution. Eustala is only
known from the Americas. Most species are
tropical, and only five species are found in
temperate North America. Another eight
tropical species have been collected in
southern Florida, southern Texas or south-

ern California. The many species in the
American tropics are probably very difficult
to separate by moi-phological characters
alone.

Key to Eustala females north of Mexico
1. Tropical species, southern California, south-
ern Texas, southern Florida ( Map 4 ) 2

- Temperate species (Map 5) 9

100 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

2(1) Scape of epigynum without annulations

(Figs. 118, 128) 3

- Scape with annulations (Figs. 138, 149) — - 4
3(2) Epigynum wider than long in both ventral

and posterior view (Figs. 118, 119);
lightly sclerotized; Texas to Panama, West
Indies devia

- Epigynum as wide as long in ventral view

(Fig. 128), longer than wide in posterior
view (Fig. 129); sclerotized; Florida,

West Indies cazieri

4(2) Middle piece of epigynum wide and large,
abnost hiding framing parts to the sides
(Figs. 167, 168); Texas to Costa Rica
- bifida

- Middle piece of epigynum narrower (Figs.

139, 150, 160) 5

5(4) Abdomen setae dilated at base; eye region
projecting slightly (Fig. 163); scape of
epigynum in side view unusually deep
(Fig. 161); Texas to Guatemala — _ clavispina

- Abdomen setae not modified, eye region not

projecting; scape of epigynum not deep

(Figs. 140, 151) 6

6(5) Epigynum in posterior view with lateral

constrictions ventrally (Figs. 139, 150) -- 7

- Epigynum without such lateral constrictions

(Figs. 180, 206); if constricted, constric-
tion dorsally (Fig. 193) 8

7(6) Epigynum in posterior view with lateral
pieces dorsally expanded, middle piece
wide ventrally (Fig. 139); abdomen with
one hump; California, Mexico califomiensis

- Epigynum in posterior view with lateral

pieces not so expanded (Fig. 150), middle
piece narrow ( Fig. 149 ) ; abdomen with
two or three large humps (Figs. 153,

154); Texas brevispina

8(6) Scape thick, finger-shaped with rounded
tip (Figs. 179, 181); epigynum in poste-
rior view long and middle piece small
(Fig. 180); tropical Florida, West Indies
eleuthera

- Scape tapering to a point (Figs. 192, 205,

228); epigynum in posterior view more or
less square in outline with middle piece
larger (Figs. 193, 206, 234, 254); whole

region 9

9(1) Epigynum in posterior view with dorsal,
lateral lobes (Fig. 193); abdomen witli
three humps in a row (Figs. 196, 197);
California to New Mexico rosae

- Epigynum in posterior view without the dor-

sal lobes ( Figs. 206, 234, 254 ) 10

10(9) Middle piece of epigynum in posterior
view larger than each lateral area (Fig.
270) and abdomen with a distinct hump
(Figs. 273, 274); California, Arizona,
northwestern Mexico conchJea

- Middle piece of epigynum smaller or as large

as lateral area ( Figs. 206, 244 ) ; if middle
piece of epigynum in posterior view larger
than lateral area, abdomen without hump;
eastern and central United States and

Canada - 11

11(10) Abdomen longer than wide with a distinct
posterodorsal hump (not in Florida)
(Figs. 209, 210); posterior median piece
of epigynum smaller in area tlian either
lateral one (Figs. 206, 280-285); total
length 5.7 to 10.0 mm, of southern Florida
specimens 5.4 mm, 0.36 (Florida), 0.44
to 0.58 mm wide anoiiera

- Abdomen, if longer than wide, posterior

tubercle indistinct (Figs. 257, 258), or
almost as wide as long (Figs. 237-241);
area of posterior median piece of epigy-
num of same size or larger than either
lateral one (Figs. 286-295); total length
less than 7.6 mm, epigynum less than 0.5

mm wide 12

12(11) Abdomen egg-shaped, longer than wide,
without tubercle, with pattern as in Figs.
258, 260; middle piece of epigynum in
posterior view distinctly larger than each
lateral one (Figs. 254, 262, 291-295).
Total length 3.4 to 7.6 mm; southern
Florida specimens 5.0 to 6.1 mm; epigy-
num 0.35 to 0.5 mm wide cfneiioni

- Abdomen almost as wide as long, subtriangu-

lar, with small posterior dorsal tubercle,
with variable pattern (Figs. 237-241);
middle piece of epigynum about the same
area as lateral ones or slightly larger; epig-
ynum with a minute posterodorsal sclero-
tized scale (Figs. 234, 244-247, 286-
290); total length 3.4 to 7.9 mm; southern
Florida specimens the smallest; epigynum
0.28 to 0.38 mm wide cepina

Key to Eustala males north of Mexico

1. Conductor of palpus very large with a tail
(Figs. 232, 250, 266, 297-312); tem-
perate and tropical 2

- Conductor of palpus without a tail and

usually small (Figs. 126-127, 136-1.37,
147-148, 157-158, 165, 174, 187, 190);
subtropical (southern Florida, southern
Texas, southern California only. Map 1 ) 7
2(1) Second femur with a ventral row of
macrosetae or at least one niacroseta
(Fig. 214); whole region anastera

- Second femur never with a ventral macro-

seta (Figs. 201, 249, 265) 3

3(2) Pacific states and southwestern states

(Map 5) 4

- Eastern states and central states (Map 5) 5

Cyclosa, Metazygia and Eustala • Levi

101

4(3) Temiinal apophysis shorter than bubble-
hke subteniiinal apophysis (Fig. 277),
conductor very large with a tail more
than twice as long as visible part of eni-
bolns (Figs. 277, 312, 318) conchlea

- Terminal apophysis overhanging bubble-

like subterminal apophysis (Fig. 202);
conductor small with tail, equal in length
to visible part of embolus (Figs. 202,

297, 313) rosae

5(3) Conductor with tail shorter than embolus
height (Figs. 215, 298^302, 314-315);
total length 3.9 to 9.5 mm, Florida males

smallest; palpus 0.9 to 1.6 mm wide

anastera

- Conductor with its tail as long or longer

than embolus height (Figs. 303-311).
Total length less than 5.0 mm; palpus

less than 1.2 mm wide 6

6(5) Terminal apophysis shorter than bubble-
like subterminal apophysis ( Figs. 266,
309-311, 317); conductor bulging
"above" embolus and with a tail about
5 times as long as wide and much longer
than embolus is high (Figs. 309-311).
Total length 3.8 to 5.0 mm; palpus 0.8
to 1.2 mm wide emertoni

- Terminal apophysis as long or longer than

bubble-like subterminal apophysis, oxer-
hanging it (Figs. 250, 303-308, 316);
conductor not bulging "above" embolus,
with a tail less tlian 4 times as long as
wide (Figs. 303-308), equal in length
or slightly longer than embolus height
(Figs. 303-308). Total length 2.5 to 4.3
mm; southern Florida specimens smallest;

palpus 0.65 to 0.72 mm wide cepina

7(1) Palpus with bubble-like semitransparent
subterminal apophysis below stylet-
shaped tenninal apophysis (Figs. 165,
174, 187, 190) 11

- Palpus lacking bubble-like semitransparent

subtemiinal apophysis or if present, distal
to ("above") stylet-shaped tenninal
apophysis (Figs. 126, 136, 147, 157) ._.. 8
8(7) Second femur without \entral row of
macrosetae (Fig. 146); palpus as in Fig-
ures 147, 148; soutliern California, Mex-
ico californiensis

- Second femur with a ventral row of macro-

setae (Figs. 125, 135, 156); palpus not
as in Fig. 147; southern Texas and Flor-
ida 9

9(8) Terminal apophysis covering embolus,
subterminal apophysis distal in palpus
(Figs. 126, 127); Texas to Panama and
West Indies devia

- Subterminal apophysis not distal and tenni-

nal apophysis not hiding embolus (Figs.
136, 157) 10

10(9) Terminal apophysis of palpus a non-
transparent lobe overhanging embolus
(Fig. 136); Florida, West Indies __.. cazieri

- Tenninal division of palpus as in Fig. 157;

Texas brevispina

11(7) Embolus twisted with embolus base ex-
tending beyond tip ( Fig. 174 ) ; Texas to
Costa Rica bifida

- Embolus hook -shaped (Figs. 165, 187,

190 ) 12

12(11) Terminal apophysis with a constricted
neck, and knife-blade-shaped tip (Fig.
165); conductor large (Figs. 165, 166);
Texas to Guatemala clavispina

- Temiinal apophysis otherwise (Figs. 187,

190); conductor small (Figs. 187, 190) ._ 13
13(12) Embolus partly liidden by bubble-like
subterminal apophysis (Fig. 190); Texas
cameronensis

- Embolus below bubble-like subtemiinal

apophysis (Fig. 187); Florida ..„ eleuthera

Eustala devia (Gertsch and Mulaik),
new combination

Figures 118-127, Map 4

NeosconeUa devia Gertsch and Mulaik, 1936, Anier.
Mus. Novitates, no. 863: 16, fig. 38, 9. Female
holotype from Edinburg, Texas, in the American
Museimi of Natural History, examined.

Eustala minima Chickering, 1955, Bull. Mus. Conip.
Zool. 112: 471, figs. 94-96, 9. Female holotype
from Barro Colorado Island, Panama Canal Zone,
in the Museum of Comparative Zoologv, ex-
amined. NEW SYNONYMY.

Note. The epigynum of specimens of E.
minima from Panama differs some from that
of specimens from the Bahama Islands and
the holotype of E. devia (Figs. 118, 119).

Description. Female holotype: Carapace
yellowish with some black patches. Poste-
rior median eye area black and lateral eyes
on black spots. Sternnm yellow, legs yel-
low with black patches and rings. Dorsum
of abdomen light and witli folium. Venter
of abdomen with white pigment spots be-
hind epigynum and a gray trapezoid and a
gray transverse band in front of spinnerets
(Fig. 124). The abdomen is almost as wide
as long and without humps. Female from
South Bimini: Total length 3.6 mm. Cara-
pace 1.5 mm long, 1.2 wide. First fepiur^

102 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

2.0 mm; patella and tibia, 2.3; metatarsus, after Prof. M. A. Cazier, collector of many

1.5; tarsus, 0.6. Second patella and tibia, specimens of tliis species in South Bimini.

1.9; third, 1.1; fourth, 1.6. Description. Female from Bimini: Cara-

Male from South Bimini: Total length 2.8 pace orange-brown with paired black
mm. Carapace 1.4 mm long, 1.2 wide. First patches and white down. Legs orange-
femur, 2.1 mm; patella and tibia, 2.4; meta- brown, indistinctly banded. Dorsum of ab-
tarsus, 1.6; tarsus, 0.7. Second patella and domen with lines oudining the folium and
tibia, 1.8; third, 0.8; fomth, 1.5. sometimes with a black longitudinal band

The illustrations were made from the f e- ( Fig. 132 ) . Venter with little black pigment

male holotype and from a South Bimini (Fig. 134). Thoracic depression a median

male. longitudinal line. Posterior median eyes

Diagnosis. The abdomen of the female 0.9 diameter of anterior, laterals 0.8 diam-

lacks a distinct hump (Figs. 122, 123). As in eter. Anterior median eyes 1.5 diameters

E. cazieri, the epigynum has a smooth scape apart, posterior median eyes 1.5 diameters

without annuli, but unlike that of E. cazieri, apart. The abdomen is triangular, pointed

the scape is tipped by a knob (Figs. 118, above spinnerets. Total length 5.4 mm.

120). The male differs from other species Carapace 2.2 mm long, 1.9 wide. First

of Eustala in having the terminal apophysis femur, 2.5 mm; patella and tibia, 3.0; meta-

covering the embolus in mesal view (Fig. tarsus, 1.9; tarsus, 0.7. Second patella and

126), and the subterminal apophysis apical, tibia, 3.0 mm; third, 1.4; fourth, 2.5.

The similar West Indian E. percUta Bryant Male from Miami: Coloration like that of

has a differently shaped terminal apophysis, female. Eye sizes about as in female, ante-

Distrihution. Southern Texas to Panama, "or median eyes slightly larger. Anterior

Bahamas, Hispaniola, Puerto Rico ( Map 4). median eyes their diameter apart, posterior

Records. Mexico. Tabasco. 2 mi. NE of median eyes slightly more than tlieir diam-

Comalcalco, S. Panama. Boquete; Arrai- eter apart. First coxa with a hook. Total

jan; El Valle; Porto Bello; all ? 2 . Cajial length 4.7 mm. Carapace 2.4 mm long, 1.9

Zone. Barro Colorado Island; Ft. Randolph; wide. First femur, 3.6 mm; patella and

Chilibre; Madden Dam; Forest Reserve; all tibia, 4.3; metatarsus, 2.8; tarsus, 1.1. Sec-

9 9. Bahamas. South Bimini, 9, ^. Haiti, ond patella and tibia, 3.0 mm; third, 1.5;

Kenskoff, 9 . Puerto Rico. Mayagiiez; Cam- fourth, 2.4.

balche Forest east of Arecibo. Female illusti-ated came from South

Bimini, male from Miami.

Eustala cazieri new species Variation. Females may lack a pattern

Figures 128-137 Mao 4 ^" ^^^^ dorsum of the abdomen, and some

have a median longitudinal dark band. To-

Holotype. Female from Plantation Key, tal length of Florida females, 5.2 to 6.8 mm,

Momoe County, 4 miles south of Tavernier, carapace 2.0 to 2.5 long, 1.8 to 2.0 wide.

Florida, 11 March 1963 (H. and L. Levi), Males vary in total length 3.3 to 4.7 mm,

edge of hardwood forest, in the Museum of carapace 2.1 to 2.4 long, 1.7 to 1.9 wide.

Comparative Zoology. The species is named Diagnosis. Females differ from other

Figures 118-127. Eustala devia (Gertsch and Mulaik): 118-121. Epigynum: 118. Ventral. 119. Posterior.

120. Lateral. 121. Posterior, cleared. 122. Female carapace and abdomen, dorsal. 123. Female, legs re-
moved, lateral. 124. Female abdomen, ventral. 125. Male, ventral macrosetae on left femora. 126, 127. Left
male palpus: 126. Mesal. 127. Ventral.

Figures 128-137. Eustala cazieri n. sp.: 128-131. Epigynum: 128. Ventral. 129. Posterior. 130. Lateral.

131. Posterior, cleared. 132. Female, dorsal. 133. Female, legs removed, lateral. 134. Female abdomen,
ventral. 135. Male, ventral macrosetae of left femora. 136, 137. Male palpus: 136. Mesal. 137. Ventral.

Cyclosa, Metazygia and Eustala ' Levi 103

Scale lines. 0.1 mm except Figs. 122-125 and 132-135, 1.0 mm.

104 BuUetin Museum of Comparative Zoology, Vol. 148, No. 3

Florida species by tlie angular abdomen
(Figs. 132, 133) and by lacking annuli on
the smooth, anteriorly directed scape of the
epigynum (Figs. 128-130). The scape lacks
the knob present in E. devia. Unlike males
of most Eusfala species, those of E. cazieri
have no bubble-like transparent subterminal
apophysis (Fig. 136); they have an ovoid
terminal apophysis overhanging tlie em-
bolus (Figs. 136', 137).

Distribution. Southern Florida and Ba-
hama Islands ( Map 4).

Records. Florida. Dade Co.: Miami;
Miami Beach. Monroe Co.: Tavernier. Ba-
hama Islands. North Bimini; South Bimini;
Crooked Isl.; Eleuthera; Great Abaco IsL;
North Caicos Isl.; Berry Isl.; Andros IsL;
New Providence.

Eustala californiensis (Keyserling),
new combination

Figures 138-148, Map 4

Cyiiophora californiensis Keyserling, 1885, Vei-
handl. Zool. Bot. Ges. Wien, 34: 525, pi. 13,
fig. 24, ? . Female holotype from "California"
in the Museum of Comparative Zoology, ex-
amined. Keyserling, 1893, Spinnen Amerikas, 4:
263, pi. 13, fig. 196, 9 . Roewer, 1942, Katalog
der Araneae, 1: 751. Bonnet, 1956, Bibliographia
Araneorum, 2(2) : 1361.

Araneus diegensis Schenkel, 1950, Verb. Natiuf.
Gesell, Basel, 61: 67, fig. 23, 9. Female holo-
type from Missions Bay, San Diego, California,
in the Natural Histoiy Museum, Basel, examined.
NEW SYNONYMY.'

Eustala abdita Chickering, 1955, Bull. Mus. Comp.
Zool., 112: 410, figs. 19-23, c5 . Male holotype
from Huajuapan, Oaxaca, Mexico in the Amer-
ican Museum of Natural History, examined.
NEW SYNONYMY.

Eustala mcxicana Chickering, 1955, Bull. Mus.
Comp. Zool., 112: 465, figs. 88-89, 9. Female

holotype from Lo Bajo, Guerrera, Mexico in the
American Museum of Natmal History, examined.
NEW SYNONYMY.

Description. Female from Oaxaca: Cara-
pace light brown with paired dark brown
patches and dark longitudinal mark in tho-
racic cleft. Legs light brown with some
black rings, more distinct ventrally. Dor-
sum of abdomen whitish with folium (Fig.
142). Sides with gray lines. The abdomen
is triangular, narrow with a dorsal posterior
hump (Figs. 142-144). Total length 5.4
mm. Carapace 2.2 mm long, 2.0 wide.
First femur, 3.2 mm; patella and tibia, 3.7;
metatarsus, 2.0; tarsus, 0.9. Second patella
and tibia, 3.0 mm; third, 1.5; fourth, 2.6.

Male from Oaxaca: Coloration as in fe-
male. Total length 3.6 mm. Carapace 1.8
mm long, 1.5 wide. First femur, 2.9 mm;
patella and tibia, 3.0; metatarsus, 1.9; tarsus,
0.9. Second patella and tibia, 2.0 mm; third,
1.1; fourth, 1.9.

Female illustrated was from Oaxaca and
males from Colima and Veracruz.

Variation. Females usually have the ab-
domen narrow (Fig. 142), but it may be
wider and may have a dorsal triangular
dark mark (Fig. 143). Total length of fe-
males is 3.9 to 7.0 mm, carapace 1.6 to 2.6
long, 1.4 to 2.1 wide. Total length of males
3.0 to 4.3 mm, carapace 1.6 to 2.3 long, 1.3
to 1.9 wide.

Diagnosis. Females can be separated
from all related species by the posterior
view of the epigynum, which shows a ven-
tral constriction with laterally expanded
dorsolateral lobes on each side (Fig. 139).
The tenuinal apophysis of the palpus ( Figs.
147, 148), lacking the usual sclerotized

Figures 138-148. Eustala californiensis (Keyserling): 138-141. Epigynum: 138. Ventral. 139, 141. Poste-
rior. 140. Lateral. 141. Cleared. 142. Female, dorsal (Mexico). 143. Female abdomen, dorsal (Califor-
nia). 144. Female, legs removed, lateral. 145. Female abdomen, ventral. 146. Male, ventral macrosetae
on left femora. 147, 148. Male left palpus: 147. Mesal. 148. Ventral.

Figures 149-158. Eustala brevispina Gertsch and Davis: 149-152. Epigynum: 149. Ventral. 150. Posterior.
151. Lateral. 152. Posterior, cleared. 153. Female carapace and abdomen. 154. Female, legs removed,
lateral. 155. Female abdomen, ventral. 156. Male, ventral macrosetae of left femora. 157, 158. Male pal-
pus: 157. Mesal. 158. Ventral.

Scale lines. 0.1 mm except Figs. 142-146, 153-156, 1.0 mm

Cyclosa, Metazygia and Eustala • Levi 105

106 Bulletin Museum of Comparative Zoology, Vol 148, No. 3

prong, is distinct from that of related spe-
cies.

Distribution. Southern California, San
Luis Potosi south to Chiapas (Map 4).

Eustala brevispina Gertsch and Davis
Figures 149-158, Map 4

Eustala brevispina Gertsch and Davis, 1936, Amer.
Mus. Novitates, 881: 12, figs. 9, 10, ?, $. Male
holotype from Cameron Co., Texas in the Amer-
ican Museum of Natural History, examined.

Description. Female: Carapace brown
with black marks and white setae. Legs
brown with narrow black bands on distal
articles. First, second and fourth femora
mostly black. The abdomen is contrastingly
marked and has two posterior humps (Figs.
153, 154). Total length 8.5 mm. Carapace
2.7 mm long, 2.4 wide. First femur, 3.2 mm;
patella and tibia, 3.9; metatarsus, 2.5; tar-
sus, 1.0. Second patella and tibia, 3.5 mm;
third, 1.7; fourth, 2.9.

Male holotype: Total length 5.8 mm.
Carapace 3.0 mm long, 2.6 wide. First
femur, 4.1 mm; patella and tibia, 4.7; meta-
tarsus, 3.0; tarsus, 1.2. Second patella and
tibia, 4.0 mm; third, 2.1; fourth, 3.6.

Diagnosis. This large species can be told
from related species by the large abdominal
humps (Figs. 153, 154) not present in E.
calif orniensis. Like E. caUforniemis the
epigynum in posterior view has a diagnostic
constriction (Fig. 150), but the lateral
pieces are differently shaped from those of
£. caUforniemis. Unlike most Eustala spe-
cies the male lacks the transparent subter-
minal apophysis. The shape of the terminal
apophysis of tlie palpus (Figs. 157, 158) is
unlike tliat of related species.

Distribution. Texas. Cameron Co.:
Brownsville, 25 May 1934, 9 allotype; 1
June 1934, 3$; 8 June 1934, 39 (J. N.

Knull). Tamaulipas. La Pesca, 17 May
1952, 19 (W.J. Gertsch).

Eustala clavispina (O.P.-Cambridge)
Figures 159-166, 176-177, Map 4

Epeira clavispina O.P.-Cambridge, 1889, Biologia
Centrali-Americana, Araneidea, 1: 37, pi. 7, fig.
11, 9. Two female syntypes from Cubilguitz in
Vera Paz, Guatemala, in the British Museum,
Natural History, examined. Keyserling, 1892,
Spinnen Amerikas, 4: 102, pi. 5, fig. 75, 9.

Amamra nigromaculata O.P.-Cambridge, 1895,
Biologia Centrali-Americana, Araneidea, 1: 155,
pi. 19, fig. 5. Female holotype from Teapa,
Tabasco, Mexico in the British Museum, Natural
History, examined.

Eustala clavispina, — F.P.-Cambridge, 1904, Bio-
logia Centrali-Americana, Araneidea, 2: 509, pi.
48, fig. 19, 9 . Roewer, 1942, Katalog der Ara-
neae, 1: 764. Chickering, 1955, Bull. Mus. Comp.
Zool., 112: 428, figs. 45-48, 9.

Eustala rosae, — Gertsch and Davis, 1936, Amer.
Mus. Novitates, no. 881: 14, fig. 11, 12, $, $.
Not E. rosae Chamberlin and Ivie.

Note. This name is not a synonym of E.
conchlea McCook as thought by Bonnet
(1955, Bibliographia Araneorum, 2(2):
1839).

Description. Female from Texas: Cara-
pace brown witli dark brownish black Y
(Fig. 163). Posterior median eyes sur-
rounded by black. Sternum maculated with
black and white pigment. Legs with con-
trasting bands on femora, spots and dark
patches on distal articles. Dorsum of abdo-
men with indistinct folium (Fig. 163). Me-
dian eye area of carapace swollen. There
are lateral abdominal humps, the second
pair indistinct, and three pairs of posterior
humps in a row (Figs. 163-164). Total
length 11.5 mm. Carapace 4.0 mm long, 3.0
wide. First femur, 4.7 mm; patella and tibia,
6.0; metatarsus, 3.9; tarsus, 1.3. Second pa-
tella and tibia, 5.3 mm; third, 2.6; fourth,
4.6.

Figures 159-166. Eustala clavispina (O.P.-Cambridge): 159-162. Epigynum: 159. Ventral. 160. Posterior. 161.
Lateral. 162. Posterior, cleared. 163. Female carapace and abdomen. 164. Female abdomen, lateral. 165,
166. Male left palpus: 165. Mesal. 166. Ventral.

Cyclosa, Metazygia and Bust ALA • Levi 107

Figures 167-175. Eustala bifida F.P.-Cambridge: 167-170. Epigynum: 167. Ventral. 168. Posterior. 169. Lat-
eral. 170. Posterior, cleared. 171. Female carapace and abdomen. 172. Female, legs removed, lateral. 173.
Female abdomen, ventral. 174, 175. Male palpus: 174. Mesal. 175. Ventral.

Scale lines. 0.1 mm except Figs. 163, 164, 171-173, 1.0 mm.

108 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

Male from Texas: Coloration like that of and the terminal apophysis is kitchen-knife-
female. Carapace with two setae within blade-shaped (Fig. 165).
median eye quadrangle. The abdomen has Distribution. Southern Texas to Guate-
some strong macrosetae on dorsum and mala (Map 4).

there are t\vo posterior humps on the tri- Records. Texas. Hidalgo Co.: 7 mi. E.

angular abdomen only. Total length 6.7 of Edinburg; Edinburg. Cameron Co.: Ran-

mm. Carapace 3.1 mm long, 2.5 wide, gerville. Mexico. San Luis Potosi. Tama-

First femur, 4.4 mm; patella and tibia, 5.8; zunchale. Veracruz. Ceno Azul. Tabasco.

metatarsus, 4.0; tarsus, 1.4. Second patella Teapa. Guatemala. Vera Paz. Cubilguitz.
and tibia, 4.3 mm; third, 2.2; fourth, 4.0.

Specimens illustrated came from Texas. Eustala bifida F.P.-Cambridge

Variation. The leg banding is less dis- Figures 167-175, 178, Map 4

tinct in some specimens. Some lack the Eustala bifida F.P.-Cambridge, 1904, Biologia

characteristic basally dilate macrosetae on Centrali-Americana, Araneidea, 2: 507, pi. 48,

the abdomen; perhaps they were broken off ^^^'- 9' 1^, $ , S ■ Female, male syntx-pes from

1, . . t., '- . . ^ San Jose, Costa Rica m the British Museum,

m collectmg. The eye region projects more Natural History, examined. Roewer, 1942, Kata-

in southern specmiens, little m northern log der Araneae, 1: 764. Chickering, 1955, Bull,

ones. Specimens from Teapa, Mexico have Mus. Comp. Zool., 112: 421, figs. 35-40, 9, $.

paired black patches on the dorsum of the Bonnet, 1956, Bibliographia Araneorum, 2(2):
abdomen, and have indications of a dorsal

fold on the posterior side of the epigynum, Description. Female syntype: Carapace

as in E. rosae. Total length of females 8.4 brown, sternum brown with white pigment

to 11.5 mm, carapace 3.4 to 4.0 long, 2.6 to in center. Legs brown, banded with black-

3.1 wide. Total length of males 6.7 to 7.3 ish brown. Dorsum of abdomen with in-

mm, carapace 3.1 to 3.7 long, 2.5 to 2.7 distinct folium, black and gray marks (Fig.

wide. 171 ) . Venter black between epigynum and

Diagnosis. This is the only Eustala spe- spinnerets with a median white longitudinal

cies of the area having a projecting eye re- line through the center, widest anteriorly,

gion (Fig. 163) and basally expanded setae fading out behind (Fig. 173). Abdomen

on the abdomen. ( The setae may be broken triangular with two posterior humps in a

off and the eye region projects only little row (Figs. 171, 172). Total length 9.0 mm.

in northern specimens.) Unlike all other Carapace 4.0 mm long, 3.1 wide. First

species, the scape of the epigynum appears femur, 4.2 mm; patella and tibia, 5.8; meta-

laterally compressed, thus deeper than wide tarsus, 3.6; tarsus, 1.3. Second patella and

(Fig. 161). The embolus of the palpus is tibia, 5.0 mm; third, 2.5; fourth, 4.4.

partly hidden by the subterminal apophysis Male syntype: Coloration like that of fe-

Figures 176, 177. Eustala clavispina (O.P.-Cambridge): 176. Male, ventral macrosetae of left femora. 177.
Female abdomen, ventral.

Figure 178. Eustala bifida F.P.-Cambridge. Male ventral macrosetae.

Figures 179-188. Eustala eleuthera n. sp. 179-182: Epigynum: 179. Ventral. 180. Posterior. 181. Lateral.
182. Posterior, cleared. 183. Female carapace and abdomen. 184. Female, legs removed, lateral. 185. Fe-
male abdomen, ventral. 186. Male, ventral macrosetae of left femora. 187, 188. Male left palpus: 187.
Mesal. 188. Ventral.

Figures 189-191. Eustala cameronensis Gertsch and Davis, male: 189. Ventral macrosetae of left femora.
190, 191. Palpus: 190. Mesal. 191. Ventral.

Scale lines. 0.1 mm except Figs. 176-178, 183-186, 189, 1.0 mm.

Cyclosa, Metazygia and Bust ALA • Levi 109

110 BitUetin Museum of Comparative Zoology, Vol. 148, No. 3

male. Total length 6.3 mm. Carapace 3.4
mm long, 2.8 wide. First femur, 4.5 mm;
patella and tibia, 5.5; metatarsus, 3.7. Third
patella and tibia, 2.2; fourth, 3.9.

The illustrated specimens came from
Costa Rica except Fig. 175 from Panama.

Diagnosis. The venter of the abdomen
of females is more contrastingly colored
than that of other species; it has a white
mark framed by black (Fig. 173). Unlike
all other species north of Mexico, E. bifida
has the middle piece of epigynum bulging
in both \cntral and posterior view (Figs.
167, 168); the lateral areas are about as
long as wide (Fig. 168). Unlike all other
species north of Mexico, E. bifida has the
embolus of the male palpus different; it is
twisted (Fig. 174) and the teraiinal apophy-
sis is a long prong widest near the tip and
extending to the outer edge of the conduc-
tor (Fig. 174).

Distribution. Southern Texas to Costa
Rica (Map 4).

Records. Texas. Cameron Co.: most
southern Palm Grove, 16 Feb. 1941, 9 (L.
I. Davis). Veracruz. Rio Blanco, 6 Nov.
1957, 9 (R. Dreisbach). Costa Rica. La
Verbena, 2 9 ( Tristan ) .

Eustala eleuthera new species
Figures 179-188, IViap 4

Holoti/pe. Male from Cape Sable, Mon-
roe County, Florida, 4 April 1958, H. V.
Weems, collector, in the Museum of Com-
parati\'e Zoology. The specific name is a
noun in apposition after tlie Bahamian Is-
land Eleuthera.

Description. Female from South Bimini:
Carapace yellow \vith white hairs in ce-
phalic region. Sternum, legs yellow. Dor-
sum of abdomen speckled with black marks.

There is an outline of folium, sometimes a
black line ( Fig. 183 ) . Venter with a central
white spot, longer than wide, black on each
side; spinnerets dark brown (Fig. 185).
Posterior median eyes 1.2 diameters of an-
terior medians, laterals 0.6 diameter of ante-
rior median eyes. Anterior median eyes 1.8
diameters apart, posterior medians their
diameter apart. The abdomen is triangular
with a pointed posterior dorsal hump, and
a second smaller hump between the dorsal
hump and spinnerets (Figs. 183, 184). Total
length 6.3 mm. Carapace 2,2 mm long, 2.0
wide. First femur, 2.7 mm; patella and
tibia, 3.4; metatarsus, 2.0; tarsus, 0.9. Sec-
ond patella and tibia, 2.7 mm; thnd, 1.5;
fourth, 2.5.

Male: Coloration slightly darker than in
female. Posterior median eyes 0.8 diameter
of anterior medians, anterior laterals 0.7,
posterior laterals 0.6 diameters. Anterior
median eyes 1.3 diameters apart, posterior
median eyes their diameter apart. Total
length 3.4 mm. Carapace 1.9 mm long, 1.6
wide. First femur, 2.5 mm; patella and
tibia, 3.0; metatarsus, 2.0; tarsus, 0.7. Sec-
ond patella and tibia, 2.2 mm; tliird, 1.2;
fourth, 1.9.

Specimens illustrated came from South
Bimini.

Variation. Some individual females are
much darker than others. Total lengtli of
females 4.1 to 6.3 mm, carapace 1.7 to 2.2
long, 1.5 to 2.0 wide. Total length of males
2.7 to 4.0 mm, carapace 1.5 to 2.1 long, 1.2
to 1.6 wide.

Diagnosis. Many females can be sepa-
rated from other species by the speckled ab-
domen (Fig. 183) and by tlie short, finger-
shaped scape of the epigynum (Fig. 179).
Unlike that of E. brevispina, the epigynum

Figures 192-204. Eustala rosae Chamberlin and Ivie: 192-198. Female: 192-195. Epigynum: 192. Ventral.
193. Posterior. 194. Lateral. 195. Posterior, cleared. 196. Lateral. 197. Dorsal. 198. Abdomen, ventral.
199-204. Male: 199. Lateral, legs removed. 200. Dorsal. 201. Ventral macrosetae on left femora. 202-204.
Left palpus: 202. Mesal. 203. Apical. 204. Ventral.

Scale lines. 0.1 mm except Figs. 196-201, 1.0 mm.

Cyclosa, Metazygia and Eustala 'Levi 111

112 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

of E. eleuthera lacks a constriction in poste- wider neck, and the conductor is of very

rior view (Fig. 180), and the median piece different shape (Figs. 190, 191). The fe-

is relatively small (Fig. 180). The male has male is unknown.

a unique, bent, half -spear-shaped terminal Record. Texas. Hidalgo Co.: 7 mi. E.

apophysis (Fig. 187) and a very small con- Edinburg, 3 Sept. 1953, S (S. Mulaik).
ductor with a proximally facing pocket in

lateral view (Fig. 188). Eustala rosae Chamberlin and Ivie

Distribution. Southern Florida, Bahamas, Figures 192-204, 297, 313, Map 5

Jamaica (Map 4). Emtala wsae Chamberlin and Ivie, 1935, Bull.

Records: Florida. Monroe Co.: 2 mi. Univ. Utah, biol sen, 2(8): 22, fig. 124, ab-

north of Flamingo; Cape Sable. Bahama Is- domen. Juvenile female holotype from Roose-

lands. South Bimini; Long Island; Crooked ^^^* ^^''' ^"^°"^' '°''-

Island; New Providence. Jamaica. St. An- Description. Female from Trinity

drew Parish: Hope Gardens. St. Thomas County, California: Carapace yellow-brown

Parish: Holland Bay. St. Ann Parish: Clare- with tiny dark dots. Sternum brown with

mont. black spots and some irregular white spots.

Legs conti'astingly banded, black on brown.

Eustala cameronensis Gertsch and Dorsum of abdomen white, brown and

Davis black with a distinct folium and a median

Figures 189-191, Map 4 longitudinal dark line from anterior to pos-

Eustala cameronensis Gertsch and Davis, 1936, terior ( Fig. 197) . Sides with tllin longitudi-

Amer. Mus. Novitates, 881: 13, fig. 13, i. Male nal black lines. The abdomen is triangular,

holotype from Cameron Co., Texas in the Amer- pointed behind, with a hump in front of tlie

ican Museum of Natural History, examined. ^^-^^^ .^^^^ ^ j^^^^^^p halfway between the

Description. Male holotype: Carapace point and spinnerets; three humps in a row

yellow-brown witli dark patches and some (Figs. 196, 197). Total lengdi 7.5 mm.

tiny black pigment spots posteriorly. Ster- Carapace 3.2 mm long, 2.7 wide. First fe-

num with black pigment spots. Legs yel- mur, 4.5 mm; patella and tibia, 5.2; meta-

low-brown. Third and fourth legs banded, tarsus, 3.2; tarsus, 1.4. Second patella and

the first two unhanded. Dorsum of abdo- tibia, 4.5 mm; third, 2.5; fourth, 3.7.

men with a black-bordered folium. Abdo- Male from San Diego Co., California:

men is oval with posterior hump indistinct. Carapace, legs and sternum much darker

Total length 4.0 mm. Carapace 2.0 mm than in female. Legs with indistinct light

long, 1.7 wide. First femur, 3.4 mm; patella spots. Venter of abdomen black with trans-

and tibia, 3.6; metatarsus, 2.1; tarsus, 0.8. verse colorless area beliind genital groove

Second patella and tibia, 2.4; tliird, 1.2; (Fig. 198). Abdominal humps as in female

fourth, 2.1. . (Figs. 199, 200). Dorsum of abdomen has

Diagnosis. The male palpus (Figs. 190, scattered macrosetae. Total length 4.3 mm.

191) resembles that of E. c/ouisp/jir/. As in Carapace 2.5 mm long, 1.9 wide. First

E. clavispina the embolus is partly hidden femur, 3.4 mm; patella and tibia, 4.0; meta-

by the transparent subterminal apophysis tarsus, 2.6; tarsus, 1.0. Second patella and

(Fig. 190), the terminal apophysis has a tibia, 3.0; thu'd, 1.7; fourth, 2.7.

->
Figures 205-217. Eustala anastera (Walckenaer): 205-211. Female (Pennsylvania). 205-208. Epigynum: 205.
Ventral. 206. Posterior. 207. Lateral. 208. Posterior, cleared. 209. Lateral. 210. Dorsal. 211. Abdomen,
ventral. 212-217. Male (Pennsylvania): 212. Lateral, legs removed. 213. Dorsal. 214. Ventral macrosetae
on left femora. 215-217. Left palpus: 215. Mesal. 216. Apical. 217. Ventral.

Scale lines. 0.1 mm except Figs. 209-214, 1.0 mm.

Cyclosa, Metazygia and Evstala • Levi 113

114 BuUctin Museum of Comparative Zoology, Vol. 148, No. 3

Variation. Total length of females 6.8 to
9.0 mm, carapace 2.3 to 3.6 long, 1.9 to 3.1
wide. Total length of males 5.0 to 5.9 mm,
carapace 2.6 to 3.0 long, 2.2 to 2.7 wide.

Diagnosis. Females differ from those of
related species by having three posterior
tubercles in a row on the abdomen (Fig.
196), and by the extra lateral lobe on the
base of the epigynum (Figs. 193, 195) in
posterior view. Males differ from most re-
lated Eustala species by the half-spear-
shaped tip of the terminal apophysis of the
palpus, much wider than its stalk ( Figs. 202,
203, 297, 313). Males differ from E. anas-
tera, which have a similar terminal apophy-
sis, by lacking macrosetae on the venter of
the second femur ( Fig. 201 ) .

Natural History. Specimens have been
collected from montane forest and juniper
woodland, and creosote brush scrub in Cal-
ifornia. Most mature individuals were col-
lected from April to August.

Distribution. Oregon, Utah to Baja Cali-
fornia, New Mexico and Cliihuahua (Map
5).

Eustala anastera (Walckenaer)

Plate 7, Figures 205-232, 280-285,
298-302, 314, 315, Map 5

Epeira anastera Walckenaer, 1841, Histoire Natur-
eUe des Insectes Apteres, 2: 33. Type, Abbot
manuscript, Spiders of Georgia, in the British
Museum, Natural History, drawing no. 381.
Copy of manuscript in the Museum of Compara-
tive Zoology, examined.^ McCook, 1893, Amer-
ican Spiders, 3: 172, pi. 8, figs. 1-4, 9, $.

Epeira eustala Walckenaer, 1841, Histoire Naturelle
des Insectes Apteres, 2: 37. Type, Abbot manu-
script, Spiders of Georgia, in the British Museum,
Natural History, drawing no. 119. Copy of
manuscript in the Museum of Comparative
Zoology, examined.

Epeira apotroga Walckenaer, 1841, Histoire Natur-
elle des Insectes Apteres, 2: 43. Type, Abbot
manuscript, Spiders of Georgia, in the British
Museum, Natural History, drawing no. 371.
Copy of manuscript in the Museum of Compara-
tive Zoology, examined.

Epeira spatidata Walckenaer, 1841, Histoire
Naturelle des Insectes Apteres, 2: 44. Type,

^ See footnote under CycJosa turbinata.

Abbot manuscript, Spiders of Georgia, in the
British Museum, Natural History, drawing no.
366. Copy of manuscript in the Museum of
Comparative Zoology, examined.

Epeira iUustrata Walckenaer, 1841, Histoire Natur-
elle des Insectes Apteres, 2: 45. Type, Abbot
manuscript, Spiders of Georgia, in the British
Museum, Natural History, drawing no. 186.
Copy of manuscript in the Museum of Compara-
tive Zoology, examined.

Epeira decolorata Walckenaer, 1841, Histoire
Naturelle des Insectes Apteres, 2: 49. Type, Ab-
bot manuscript. Spiders of Georgia, in the
British Museum, Natural History, drawing no.
345. Copy of manuscript in the Museiun of
Comparative Zoology, examined.

Epeira triflex Walckenaer, 1841, Histoire Naturelle
des Insectes Apteres, 2: 60. Type, Abbot manu-
script, Spiders of Georgia, in the British Museum,
Natural History, illustration no. 112. Copy of
original in the Museum of Comparative Zoology,
examined.

Epeira trinotata Walckenaer, 1841, Histoire Natur-
elle des Insectes Apteres, 2: 75. Type, Abbot
manuscript. Spiders of Georgia, in the British
Museum, Natural History, illustration no. 272.
Copy of original in the Museum of Comparative
Zoology, examined.

Eustala anastera, — Chamberhn and Ivie, 1944,
Bull. Univ. Utah, biol. ser., 7(5): 102, fig. 4.
Kaston, 1948, Bull. Connect. Geol. Nat. Hist.
Surv., 70: 233, figs. 706-709, 727, $. Bonnet,
1956, Bibliographia Araneorum, 2(2): 1837 (in
part only).

Note. I have listed only the first Abbot
figure cited by Walckenaer for each name.
Epeira circulata Walckenaer, 1841, p. 79,
may have been an Eriophora, probably not
Eustala as indicated by Chamberlin and
Ivie (1944).

Description. Female from Pennsylvania:
Carapace brown, sides of thorax darker.
Thorax covered with white hairs and down.
Legs dark, banded. Dorsum of abdomen
with folium or longitudinal dark line. Ab-
domen triangular, longer tlian wide with a
distinct posterior dorsal hump (Figs. 209,
210). Total length 7.5 mm, carapace 2.7
mm long, 2.4 wide. First femur, 3.4 mm;
patella and tibia, 4.2; metatarsus, 2.4; tarsus,
0.9. Second patella and tibia, 3.6 mm; third,
1.7; fourth, 3.2.

Male from Pennsylvania: Coloration as
in female. Total length 4.8 mm. Carapace

Cyclosa, Metazygia and Eustala • Levi 113

218

219

220

223

221

222

Figures 218-232. Eustala anastera (Walckenaer): 218-222. Dorsal patterns of female abdomen. 218. (Penn-
sylvania). 219. (West Virginia). 220. (Michigan). 221, 222. (Florida). 223, 224. Female abdomen, lateral. 223.
(Georgia). 224. (southern Texas). 225. Female eye region and chelicerae. 226-230. Epigynum. 226-228:
Ventral. 229, 230. Posterior. 226, 227. (Kerr Co., Texas). 228. (Goliad Co., Texas). 229. (Nova Scotia, Can-
ada). 230. (Cimarron Co., Oklahoma). 231. Left and right palpi from same individual (Grant Par., Louisiana).
232. Left palpus expanded.

Scale lines. 0.1 mm except Figs. 218-225, 1.0 mm.

Abbreviations, a, terminal apophysis; c, conductor; dh, distal hematodocha; e, embolus; m, median apophysis;
r, radix; sa, subterminal apophysis; t, tegulum.

116 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

2.5 mm long, 2.2 wide. First femur, 3.6 mm; Total length of females 5.4 to 10.0 mm,

patella and tibia, 4.3; metatarsus, 2.9; tarsus, carapace 2.5 to 3.3 long, 2.3 to 2.6 wide.

1.2. Second patella and tibia, 3.0 mm; third, Total length of males 3.9 to 9.5 mm, cara-

1.8; fourth, 2.8. pace 2.2 to 4.8 long, 1.8 to 4.0 wide. The

Variation. The pattern is variable al- smallest specimens all came from central

though most specimens are dark. Some and southern Florida.

specimens have a folium on the abdomen Diagnosis. The species differs from the

(Figs. 210, 218, 221), some have black related E. eniertoni and E. cepina hy being \

patches on white (Figs. 219, 222), others darker and larger. Females differ from E.

only a longitudinal median line (Fig. 220). cepina by size, color and shape of the abdo-

Females from Texas have two humps in a men (Figs. 209, 210), from E. emertoni by

line (Fig. 224), as do occasional females having the median area of the base in poste-

from other areas (Fig. 223); one from rior view smaller than each lateral area

Cimarron, Oklahoma, had three. The hump (Figs. 280-285) (but in eastern Canada

is smaller in the northeastern part of the where emertoni is absent, the epigynum may

range, where E. emertoni is not found, and resemblethat of £. emerfoni). The abdomen

the epigynum has a larger middle piece in is longer than wide with a distinct posterior

posterior view (Fig. 229) and resembles hump (unlike £. emertoni). The epigyna

that of E. emertoni. are larger, 0.36 (Florida), to 0.58 mm wide
Males sometimes have a short and (the smallest from south centi-al Florida)
pointed terminal apophysis (Fig. 315). Such than those of E. cepina. Females differ
males included those collected and deter- fiom the western E. rosae and E. conchlea
mined by W. Ivie from eastern Fennsyl- by the shape of the epigynum in posterior
vania, one each from Alabama ( Mobile ) , view. The contrasting black and white pat-
Mississippi (Jefferson City), West Virginia, tern (Figs. 219, 222) found in some indi-
Virginia, South Dakota, Idaho, Texas, Okla- viduals is diagnostic; it is not found in
homa, Mississippi, Ontario, Connecticut related species.

(New Canaan), and all males from Michi- Males differ from E. rosae, E. emertoni
gan. The smaller males in Florida may have and E. cepina by having a row of three to
only one macroseta on the second femur, five short macrosetae on the venter of the
sometimes only on one side. But several second femur (Fig. 214); rarely, in small
very large males lacked these macrosetae Florida specimens, there is only one. (These
entirely: one from Calhoun Co., Arkansas, macrosetae may be absent in individual
one from Boston, Mass., one from Lebanon males and always absent in those from east-
State Forest, New Jersey and one from Cen- ern Canada. ) The conductor is smaller and
ter Harbor, New York. Most males from the its tail shorter than that of E. triflex and E.
northeastern part of the range, where E. cepina (Figs. 298-302). Most of the con-
emertoni and E. cepina are not found, lack ductor is "above" the embolus, the portion
these macrosetae and the outer, "upper," "below" the embolus is less in length than
bulge of the conductor is smaller (Fig. 298). the embolus height. The terminal apophysis

Figures 233-252. Eustala cepina (Walckenaer): 233-236. Epigynum. (Pennsylvania): 233. Ventral. 234. Pos-
terior. 235. Lateral. 236. Posterior, cleared. 237-239. Dorsal patterns of female abdomen (all Texas):
240-242. Female (Pennsylvania): 240. Lateral, legs removed. 241. Dorsal. 242. Abdomen, ventral. 243-248.
Epigynum: 243, 245. Ventral. 244-248. Posterior. 243, 244. (New Jersey). 245, 246. (Missouri). 247. (Kan-
sas). 248. (Emmet Co., Michigan). 249-252. Male (Pennsylvania): 249. Ventral macrosetae of left femora.
250-252. Left palpus: 250. Mesal. 251. Apical. 252. Ventral.

Scale lines. 0.1 mm except Figs. 237-242, 249, 1.0 mm.

Cyclosa, Metazygia and Eustala • Levi 117

118 Bulletin Museum of Comparative Zoology, Vol. 148, No. 3

tip is usually (but not always) "half-spear-
shaped" and wider than its neck ( Figs. 215,
314), but not so wide as that of E. rosae. In
both E. friflex and E. cepina, the terminal
apophysis tip is only rarely wider than its
neck. In £. rosae it is wider but, as in the
other two species, lacks tlie line of ventral
macrosetae on the second femur. The con-
ductor of the palpus, unlike that of E. ce-
pina, has an "upper, outer" lobe ( seen upper
right in Figs. 299, 302); the palpus is much
larger (0.91 to 1.6 mm wide in mesal view)
than that of E. cepina.

Natural History. Eustala anastera is
commonly found as prey in Trypoxylon and
Trypargilum mud-dauber wasp nests. The
species is found in diverse habitats. Col-
lecting sites are goldenrod (Solidugo) fields,
chokeberry, an apple tiee in Ontario, a
tamarack bog {Larix occidentalls) in Mani-
toba, a balsam fir ti-ee (Abies halsamea) in
New Brunswick, a white spruce (Picea
glauca) in New Brunswick, maple woods in
Wisconsin, and tamarack (Larix occiden-
talis). Specimens have also been collected by
sweeping a marsh, in xeromesic woods, by
beating dead oak branches, by sweeping
Poa pratense, in a web in dead twig in
Michigan, in loblolly pine (Piniis taeda) in
Arkansas; in oak-pine flatwoods, by sweep-
ing tiu-key oak (Quercus laevis) scrub, by
sweeping cypress (Taxoditim) swamp edge,
in palm-cypress (Taxodium), in red man-
grove (Rhizophora) hammock, along a road,
in a web in Spanish moss ( Tillandsia iisne-
oides), on Pimis clausa, and near scrub oak
in Florida. I think one requirement for Eu-
stala anastera is dead branches in a rela-
tively open wooded area or along wood
borders. Comstock (1940) reports vertical
webs from low bushes. The spider does not
make a retreat but sits on bark or dead
branches to the side of the web, "the spider
closely resembling the bark of the tree or
other plant on which it rests; and they act
as if conscious of this protection, limning
only a short distance when disturbed and
then crouching down close to the bark."
Kaston (1948) reports Eustala anastera as

being one of the few orb-weavers oveiAvin-
tering in the penultimate instar. Males are
found throughout the season but, even in
Florida, are more common in spring. The
web illustrated by Plate 7 is described in i
the introduction as belonging to Eustala.
Distribution. Throughout southern Can-
ada and the United States, except perhaps
within the range of the similar E. rosae in
CaHfomia. The southern limits are un-
known but are believed to be Central Amer-
ica (Map 5).

Eustala cepina (Walckenaer)

Figures 233-252, 286-290, 303-308,
316, Map 5

Epeira cepina Walckenaer, 1841, Histoire Natur-
elle des Insectes Apteres, 2: 37. Type, Abbot
manuscript. Spiders of Georgia, in the British
Museum, Natural History, drawings no. 173 and
175. Copy of original in Museum of Compara-
tive Zoology, examined.

Epeira parvula Keyserling, 1863, Sitzungsber.
Naturf. Gesellsch. Isis, Dresden, p. 131, pi. 6,
figs. 9, 10, 9 . Female lectotype here designated
from Baltimore in the British Museum, Natural
History. One female, one male paralectotypes
are E. cepina, another female E. anastera; one
juvenile female paralectotype from Peoria is E.
anastera. NEW SYNONYMY.

Eustala cepina, — Chamberlin and Ivie, 1944, Bull.
Univ. Utah, biol. sen, 7(5): 103.

Eustala arkansana, — Archer, 1951, Amer. Mus.
Novitates, no. 1487: 19, fig. 47, ?. Female
allotype not male holotype.

Note. Most specimens in collections had
been labeled E. anastera, but A. F. Archer
called this species A. triflex.

Description. Female from Pennsylvania:
Carapace orange-brown. Sternum orange-
brown with some white spots. Legs orange-
brown, slightly banded. Dorsum of abdo-
men with a folium. The abdomen is
triangular, almost as wide as long (Figs.
237-241). Total length 5.5 mm. Carapace
2.3 mm long, 1.7 wide. First femur, 3.2 mm;
patella and tibia, 3.6; metatarsus, 1.9; tarsus,
0.8. Second patella and tibia, 2.6 mm;
third, 1.4; fourth, 2.2.

Male: Coloration as in female. The ab-
domen is oval, triangular. Total length 3.3

Cyclosa, Metazygia and Eustala • Levi 119

Figures 253-268. Eustala emertoni (Banks): 253-259. Female (Pennsylvania): 253-256. Epigynum: 253. Ven-
tral. 254. Posterior. 255. Lateral. 256. Posterior, cleared. 257. Lateral. 258. Dorsal. 259. Female abdomen,
ventral. 260. Dorsal pattern of female abdomen (Texas). 261-264. Epigynum (Connecticut): 261. Ventral.
262. Posterior. 263. Lateral. 264. Posterior, cleared. 265-268. Male (Pennsylvania): 265. Ventral macro-
setae of left femora. 266-268. Left palpus: 266. Mesal. 267. Apical. 268. Ventral.

Scale lines. 0.1 mm except Figs. 257-260, 265, 1.0 mm.

120 Bulletin Muscud) of Comparative Zoology, Vol. 148, No. 3

mm. Carapace 1.7 mm long, 1.5 \\dde.
Fii-st femur, 2.2 mm; patella and tibia, 2.7;
metatarsus. 1.8; tarsus, 0.8. Second patella
and tibia, 2.3 mm; third, 1.0; fourth, 1.8.

Voriation. The color \ariation of the ab-
domen is less than that of E. amistera, a
black median longitudinal line is common
(Figs. 237-241). Total length of females
3.4 to 7.9 mm, carapace 1.4 to 2.9 long, 1.3
to 2.2 wide. Total length of males 2.5 to
4.3 mm, carapace 1.5 to 2.4 long, 1.2 to 2.0
wide. The smallest specimens, females
measuring total length 3.4 to 4.5 mm, epigy-
num less than 0.38 mm wide, all came from
southern Florida. Some specimens appear
intermediate with E. einertoni and perhaps
E. a nasi era.

Diapwsis. The abdomen is triangular
(Figs. 237-241), almost as wide as long,
and ma)- lack a dorsal pattern. The middle
area of the epigynum in posterior \dew is
larger or subequal to the lateral (unlike E.
emertoni) (Figs. 236, 244, 247, 286-290).
The epig)'num is much smaller in size (0.28
to 0.38 mm wide) than that of E. armstera.
The male differs from E. emertoni in that
the longer terminal apophysis overhangs the
bubble-like subtenninal apophysis (Figs.
303-308, 316). The conductor lacks the lobe
(to the upper right in Figs. 303-308) pres-
ent in both E. anastera and E. emertoni.
The conductor is smaller (Figs. 303-308)
than that of E. emertoni and E. anastera.
The embolus sits in the middle of the con-
ductor, not in the "lower" half as is com-
mon in E. anastera. The palpus is also
always smaller in size (about 0.65 to 0.72
mm wide) than in E. anastera and E. emer-
toni.

Natural History. Eustala cepina is com-
monly found as prey in mud-dauber wasp
nests, of Chahjhina wasps in Oklahoma. It
has been found on lake shores in Michiuan
and \\dsconsin, in dune grass and mixed for-
est in \\dsconsin, by sweeping weeds in
Illinois, in pine dunes in Indiana, by sweep-
ing around a pond in Pennsylvania, in a
garden in North Carolina, on pecan trees in

South Carolina, in low grass and an urban
area in Alabama, in pond vegetation, on
wheat and cotton in Arkansas, on weeds
l)eside a road in Mississippi, by beating
cedar (Taxodium) branches on a slope
near a stream in Georgia; in oaks along a
beach, in grasslands, and on a small oak in
an open area in mesic hammock in Florida.
The spiders also probably rest on dead
branches next to tlie web without retreat.
It seems to prefer wetter areas than does
E. ana.stera.

Distribution. New England south to
Florida, Ontario, Wisconsin, Colorado,
central Texas to Mexico (Map 5).

Eustala emertoni (Banks)

Figures 253-268, 291-295, 309-311,
317, Map 5

? Epcira petasata Walckenaer, 1841, Histoire
Naturelle des Insectes Apteres, 2: 70. Type,
Abbot manuscript. Spiders of Georgia, in tlie
British Museum, Natural History, illustration no.
135. Copy of original in Museum of Compara-
tive Zoology, examined. Doubtful name.

Epeira emertoni Banks, 1904, J- New York Ento-
mol. Soc., 12: 111. Female syntypes from Sea
Cliff, N.Y., Washington, D.C., and Auburn, Ala-
bama, lost.

Eustala triflex, — Chamberlin and Ivie, 1944, Bull.
Univ. Utah, biol. ser., 8(5): 103 (not £. triflex
Walckenaer ) .

Eustala arkansana Archer, 1951, Amer. Mus. Novi-
tates, no. 1487: 19, fig. 44, $, not 9. Male
holotype from Berryville, Carroll Co., Arkansas,
in the American Museum of Natural History',
examined. NEW SYNONYMY.

Note. This species, called Eustala triflex
by Chamberlin and Ivie (1944), is not E.
triflex Walckenaer. Eustala triflex, fig. no.
112 of Abbot's manuscript. Spiders of
Georgia, is contrastingly white and black
as in Figures 219, 222, a coloration not
found in this species. The same comment
applies to the name E. trinotata Walcke-
naer, Abbot's fig. no. 272. Chamberlin and
Ivie believed Hentz's bombycinaria to be
this species. The light shoulder spots of
bombycinaria Hentz are found, as in fig. 16,
plate 31, but they are not wliite as shown by
Hentz in the figures and in a colored manu-

Cyclosa, Metazygia and Eustala ' Levi 121

Figures 269-279. Eustala conchies (McCook): 269-275. Female: 269-272. Epigynum: 269. Ventral. 270. Pos-
terior. 271. Lateral. 272. Posterior, cleared. 273. Abdomen, lateral. 274. Dorsal. 275. Abdomen, ventral.
276. Male ventral macrosetae on left femora. 277-279. Male left palpus: 277. Mesal. 278. Apical. 279. Ven-
tral.

Scale lines. 0.1 mm, except Figs. 273-276, 1.0 mm.

script illustration at Harvard University.
Banks was the first to describe tlie species
and give diagnostic characters.

Description. Female from Pennsylvania:
Head region much Hghter than sides of
thorax. Carapace with white down. Ster-
num with some black pigment marks. Legs
with onlv femora banded. Dorsum of ab-

domen with very distinct contrasting folium
(Fig. 258). The abdomen is oval without
hump (Figs. 257, 258). Total length 5.6
mm. Carapace 2.3 mm long, 2.0 wide. First
femur, 3.0 mm; patella and tibia, 3.6; meta-
tarsus, 1.9; tarsus, 0.7. Second patella and
tibia, 2.9 mm; tMrd, 1.5; fourth, 2.5.

Male from Pennsylvania: Coloration and

122 Bulletin Museum of Cojnparative Zoology, Vol. 148, No. 3

sliapc as in female. Total length 4.3 mm.
Carapace 2.3 mm long, 1.7 wide. First
femur, 3.0 mm; patella and tibia, 3.4; meta-
tarsus. 2.0; tarsus. O.S. Second patella and
tibia, 2.4; third, 1.4; fourth, 1.9.

\'(iri(ition. The abdominal pattern is
similar in most specimens (Fig. 258), some
lKi\t' ;i dark triangle on the dorsum (Fig.
260). Total length'of females 3.4 to 7.6 mm,
carapace 1.7 to 3.5 long, 1.5 to 2.9 wide.
Total length of males 3.8 to 5.0 mm, cara-
pace 2.4 to 2.6 long, 1.8 to 2.2 wide. South-
ern Florida females measure 5.0 to 6.1 mm
total length, carapace 2.0 to 2.3 long, 1.7 to
2.0 wide.

Di(iii,nosis. The abdomen of females is
egg-shaped, widest anteriorly, the posterior
hump absent; the median area of the epigy-
iium in posterior view is distinctly larger
than the small posterior lateral areas (Figs.
254, 262, 291-295), unlike that of E. anas-
terci and, usually, of the smaller E. cepina.
The epigynum is 0.4 to 0.5 mm wade, larger
than that of E. cepina. It is similar to the
epigynum of E. conchlea in the west, but
the abdominal hump of E. conchlea is lack-

ing in E. cnicrtoni.

Males lack macrosetae on the venter of
the second leg femora (Fig. 265). The tip
of the terminal apophysis of the palpus is
shorter than the bubble-like subterminal
apophysis, unhke tliat of E. cepina. The
conductor is much larger than that of E. ce-
pina and E. anasfera and, unlike that of E.
cepina, is bulging "on top" and has a thin
tail, about five times as long as wide ( Figs.

266, 309-311, 317). The palpus is 0.8 to 1.2
mm wide.

Nattiral History. Specimens have been
collected as prey by ChuIyJ)ion wasps in
Oklahoma and other mud-dauber wasps, in
button woods (Platanus sp.) in Rhode Is-
land (the northernmost locality), in wheat
and in alfalfa in Arkansas, in a broom-sedge
(Andropogon virginictis) field and bottom-
land pine-hardwood forest in North Caro-
lina, in pinewoods and salt marsh in Georgia,
in roadside low weeds and grass in Missis-
sippi, in a wooded area in Texas, in pine-
flatwoods, bottomland, palmetto flatwoods, j
and around a swamp in Florida.

Distribution. Rhode Island, Michigan to
Florida, Kansas, central Texas and north-
eastern Mexico (Map 5).

Eustala conchlea (McCook)

Figures 269-279, 296, 312, 318,
Map 5

Epcira parvula var. conchlea McCook, 1888, Pioc.
Acad. Sci. Philadelphia, p. 199, fig. 6, $ . Spec-
imens from Wisconsin and California. Female ,j
lectotype from California, here designated, and \
numerous female and male paralectotypes of |
the same species and one female paralectotype
which is E. califomiensis, all in the Academy of
Natural Sciences, Philadelphia, examined. Wis-
consin specimens do not survive.

Epcira anastcia var. conchlea McCook, 1893,
American Spiders, 3: 174, pi. 8, fig. In.

Eustala anastcia biiliafera Chamberlin, 1924, Proc.
Cahf. Acad. Sci., 4 ser., 12: 650. Female holo-
type from Isla Partida, GuLf of California in the
California Academy of Sciences, examined.
NEW SYNONYMY.

Figures 280-296. Epigyna of the E. anasfera group.

Figures 280-285. Eustala anastera (Walckenaer): 280-282. Posterior: 283-285. Mounted and cleared on a
microscope slide: 280. (Michigan). 281. (northern Florida). 282. (Texas). 283. (Pennsylvania). 284 (High-
land Co., Florida). 285. (Texas).

Figures 286-290. Eustala cepina (Walckenaer) mounted and cleared: 286. (New Jersey). 287. (Pennsylvania)
288. (Florida). 289. (Missouri). 290. (Kansas).

Figures 291-295. Eustala emertoni (Banks): 291-293. Mounted and cleared: 291. (Pennsylvania). 292. (Flor-
ida). 293. (Texas). 294, 295. Posterior: 294. (Florida). 295. (Texas).

Figure 296. Eustala conctilea (McCook).

Scale line. 0.1 mm.

Cyclosa, Metazygia and Eustala ' Levi 123

124 BuUrtin Mu.scum of Comparative Zoology, Vol. 148, No. 3

Eustala anastera h-iica Chconiberlin. 1924, Proc.
Calif. Acad. Sci., 4 ser., 12: 050. Female holo-
type in poor ph>sical condition from Santa Inez
Island, Gulf of California in the Academy of
Sciences, examined. \K^^' SYXOXYMV.

Note. In 1935 (p. 22), Chamberlin and
Ivie compared tlie new E. ro.sae to E. conch-
lea, presumably eonsidering them sympatric.
In 1944, liowexer, they consider "Epcira
unmlcra \ar. conchlea McCook, Ibid., 173
(in part, including type)" a synonym of
E. anastera.

Descri))ti(ni. Female from Laguna Beach,
California: Head region yellow-brown, tho-
racic region darker. Black rings around pos-
terior median eyes. Sternum with black
pigment. Legs banded. Dorsum of abdo-
men with folium and median longitudinal
dark line (Fig. 274). The abdomen has one
posterior dorsal hump. Total length 5.3 mm.
Carapace 2.4 mm long, 1.9 wide. First
femur, 3.3 mm; patella and tibia, 3.9; meta-
tarsus, 1.9; tarsus, 0.9. Second patella and
tibia, 3.0 mm; third, 1.6; fourth, 2.6.

Male from Los Angeles, California: Cara-
pace more evenly brown than in female and
legs less banded. The posterior dorsal
tubercle of the abdomen is distinct. Total
length 5.2 mm. Carapace 2.7 mm long, 2.2
wide. First femur, 4.1 mm; patella and
tibia, 4.5; metatarsus, 2.9; tarsus, 1.1. Sec-
ond patella and tibia, 3.5 mm; third, 1.9;
fourth, 3.0.

Variation. The variation is less than in
other species. One female had a second
tubercle below the posterior dorsal hump.
Females measure total length 4.6 to 7.9 mm,
carapace 2.2 to 3.2 mm long, 1.8 to 2.6 wide.
Males, total length 4.3 to 5.0 mm, carapace
2.2 to 2.6 long, 1.9 to 2.2 wide. Specimens
from Baja California and Arizona tend to be
larger; a male from Tucson, Arizona, 6.4 mm
long, carapace 3.4 long, 2.7 wide.

Diagnosis. The shorter, pointed terminal
apophysis of tlie palpus (Figs. 277, 278, 312,
318), the posterior view of the epigynum
(Fig. 270), and tlie single posterior dorsal
hump (Fig. 273) separate E. conchlea from
the sympatric E. rosae. Eiistala conchlea is
very similar to the eastern North American
E. emertoni. Specimens differ from E.
emertoni by having a posterior dorsal hump
on the abdomen, while the abdomen of E.
emertoni is egg-shaped, narrow behind
witliout hump.

Natural History. This species has been
collected from grassy fields, from tall weeds,
and from reeds along a lagoon.

Distribution. Central California coast,
Arizona, Baja California and Sinaloa.

REFERENCES CITED

Adanson, M. 1763. Families des plantes, Paris,

Vincent, 190 pp.
Archer, A. F. 1951. Studies in the orb-weaving

spiders (Argiopidae) 1. Amer. Mus. Novitates,

1487: 1-52.

Figures 297-312. Conductor (stippled), embolus and terminal apoptiysis tip (black) and median apophysis
(white) of left palpus of E. anastera group. (Note different enlargements.)

Figure 297. Eustala rosae Chamberlin and Ivie.

Figures 298-302. Eustala anastera (Walckenaer): 298. (Nova Scotia, Canada). 299. (Michigan). 300. (Mis-
souri). 301. (Florida). 302. (southern Texas).

Figures 303-308. Eustala cepina (Walckenaer): 303. (Massachusetts). 304. (Wisconsin). 305. (Missouri). 306.
(southern Florida). 307. (Alabama). 308. (southern Texas).

Figures 309-311. Eustala emertoni (Banks): 309. (Georgia). 310. (northeastern Texas). 311. (southern Texas).

Figure 312. Eustala conchlea (McCook).

Figures 313-318. Distal parts of palpus of the E. anastera group. (Note different enlargements.) 313. Eustala
rosae. 314, 315. Eustala anastera. 314. (New Jersey). 315. (Pennsylvania). 316. Eustala cepina (Pennsyl-
vania). 317. Eustala emertoni (Florida). 318. Eustala conchlea.

Scale lines. 0.1 mm.

Cyclosa, Metazygia and Evstala • Levi 125

126 Bulletin Muxcinn of Comparative Zoology, Vol. 148, No. 3

Beiirv, J. \\. 1970. Spiders of the North Caro-
lina Piedmont old-field communities. J. Elisha
Mitchell Sci. Soc., 86: 97-105.

Biiv.vNT, E. 19 10. Cuban spiders in the Museum
of Comparati\e Zoology. Bull. Mus. Comp.
Zool.. 86: 247-532.

CiiAMiiKHLiN, R. V. AND W. IviK. 1935. Miscel-
laneous new American spiders. Bull. Univ.
Utah, biol. ser., 2(8): 1-79.

CnA.\uiEHLix, R. V. ANO W. IviE. 1944. Spidcrs
of the Georgia Region of North America. Bull.
Univ. Utah, biol. ser., 8(5): 1-267.

CEncKERixG, A. M. 1955. The genus Fyiistala
(Araneae, Argiopidae) in Central America.
Bull. Mus. Comp. Zool., 112: 391-518.

CoMSTOCK, J- 1940. The spider book, rev. W.
J. Gertsch, Garden City.

Kaston, B. J. 1948. Spiders of Connecticut.
Bull. State Gcol. Natur. Hist. Surv., 70: 1-
874.

LuTZ, F. A. 1915. List of Greater Antillan spi-
ders with notes on their distribution. Ann.
New York Acad. Sci., 26: 71-148.

McCooK, H. C. 1887. Note on Ctjrtophora bi-

fuica (n. sp.) and her cocoons, a new orb
weaving spider. Proc. Acad. Natur. Sci., Phila-
delphia, 1887: 342-343.

Marsox, J. E. 1947. Some observations on the
variations in the camouflage devices used by
Cyclosa iusiilaua (Costa) an Asiatic spider in
its web. Proc. Zool. Soc. London, 117: 598-
605.

RoBixsoN, M. H., Y. D. Lubix and B. Robinson.
1974. Phenology, natural history and species
diversity of web-building spiders on three
transects in Wau, New Guinea. Pacific In-
sects, 16: 117-164.

Simon, E. 1900. Fauna Hawaiiensis, Clay and
Sons, London, 2(5).

Simon, E. 1928. Les arachnides de France, 4:
553-772, Paris.

WiEHLE, H. 1928. Beitrage zur Biologic der
Araneen, insbesondere zur Kenntnis des Rad-
netzbaues. Z. Morphol. Okol. Tiere, 11:
115-151.

Worth, C. B. 1940. Protective shape and color-
ation of the spider Cyclosa biftirca (Arach-
nida: Argiopidae). Ent. News, 51: 17-18.

Cyclosa, Metazygia and Eustala • Levi 127

INDEX

Valid names are printed in italics. Page numbers refer to main references, starred page numbers to il-
lustrations.

abdita, Eustala 104

accentonotata, Parazygia 83

albonigra, Metazygia 94

algciica, Cyclosa 81*

anastera, Epeira 114, 122

anastera, Eustala 68*, 113*, 114, 115*, 123*, 125*

apotroga, Epeira 114

arkansana, Eustala 118, 120

bifida, Eustala 107*, 108, 109*
hijurca, Cijclusa 66*, 86, 89*
bifurca, Cyrtophora 86
bifiucata, Epeira 84
hrevispina, Eustala 105*, 106
buliafera, Eustala anastera 122

californiensis, C>'rtophora 104

californiensis, Eustala 104, 105*

cameronensis, Eustala 109*, 112

canadensis, Epeira 78

caroli, Cyclosa 64*, 82, 85*

caroli, Epeira 82

carolinalis, Epeira 94

carolinalis, Mctazyf^ia 94, 95*

caudata, Cyclosa 80

caudata, Epeira 80

cazieri, Eustala 102, 103*

cepina, Epeira 118

cepina, Eustala 117*, 118, 123*, 125*

cla\ispina, Epeira 106

clavispina, Eustala 106, 107*, 109*

conchlea, Epeira anastera 122

conchlea, Epeira par\'ula 122

conchlea, Eustala 121*, 122, 123*, 125*

conica, Aranea 78

conica, Cyclosa 62*, 77*, 78, 79*

conigera, Cyclosa 83

cuadrituberosa, Cyclosa 85

culta, Cyclosa 80

Cyclosa 73

decolorata, Epeira 114
devia, Eustala 101, 103*
devia, Neosconella 101
diegensis, Araneus 104
dilatata, Aranea 92

eleuthera, Eustala 109*,
elongata, Cyclosa 83
emertoni, Epeira 120
emertoni, Eustala 119*,
Eustala 96

110

120, 123*, 125*

eustala, Epeira 114

fissicauda, Cyclosa 86

illustrata, Epeira 114
index, Cyclosa 80
insulana, Cyclosa 81*

keyserlingi, Metazygia 94

lacerta, Cyclosa 82

leuca, Eustala anastera 124

Metazygia 89
mexicana, Eustala 104
minima, Eustala 101

nanna, Cyclosa 80
nigromaculata, Amamra 106

oculata, Cyclosa 81*

pallidulus, Araneus 94
Parazygia 73
parvula, Epeira 118, 122
petasata, Epeira 120

wsae, Eustala 111*, 112, 125*
rosae, Eustala 106

sienae, Cyclosa 81*
spatulata, Epeira 114

trifida, Cyclosa 84
triflex, Epeira 114
triflex, Eustala 120
trinotata, Epeira 114
tuberculifera, Cyclosa 80
turhinata, Cyclosa 63*, 79^*
tiubinata, Epeira 80

SO, 83='

vanbruysselii, Singa 80

walckenaeii, Cyclosa 65*, 84, 87*
walckenaerii, Epeira 84
\\alckenaerii, Tiuckheimia 84
wittfeldae, Epeira 92
wittfeldae, Metazygia 67*, 92, 93*

zilloides, Epeira 92
zilloides, Metazygia 92, 95*

bulletin OF TH

seum

us ISSN 0027-4100

A Classification of the Tyrant Flycatchers

(Tyrannidae)

MELVIN A. TRAYLOR JR.

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 4
29 NOVEMBER 1977

PUBLICATIONS ISSUED

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SPECIAL PUBLICATIONS.

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2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini-
dae ( Mollusca: Bivalvia). 265 pp.

3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms.
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4. Eaton, R. J. E., 1974. A Flora of Concord. 236 pp.

Other Publications.

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Price list and catalog of MCZ publications may be obtained from Publications
Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 02138, U.S.A.

A CLASSIFICATION OF THE TYRANT FLYCATCHERS
(Tyrannidae)

MELVIN A. TRAYLOR JR.

CONTENTS

Abstract 129

Introduction 129

Materials and Methods — - 130

External Morphology 130

Distribution 13 1

Literature 131

Methods 133

Limits of tlie Family 133

Classification to Subfamilies 136

Sequence and Relations of Genera 145

Elaeniinae 145

Fluvicolinae 159

Tyranninae 166

Phylogeny 171

Summary 174

List of References 175

Appendix A — Sequence of genera witli

synonyms 177

Appendix B — Generic changes since

Hellmayr 178

Appendix C — Ames' groups of genera 179

Appendix D — Warter's figures and tables .— 179

Index 183

Abstract. The classification of the Tyrannidae
is re-examined, using available morphological,
anatomical, behavioral and zoogeographic data.
The usual seven subfamilies are reduced to three,
and these are defined primarily on cranial char-
acters. The former subfamily Myiarchinae is
di\ided between the Fknicolinae and Tyranninae,
and the remaining "subfamilies" are lumped into
one subfamily, the Elaeniinae. Thirty-five genera
ii'cognized in the last complete list of flycatchers
( I lellmayr, 1927 ) are synonymized, one is resur-
rected, and two new ones are recognized, of which
one is here described.

' Bird Division, Field Museum of Natural His-
tory, Roosevelt Road at Lake Shore Drive, Chi-
cago, IL 60605.

INTRODUCTION

This investigation of the classification of
the New World flycatchers, Tyrannidae,
was undertaken as the first step in the
preparation of a list of the family for in-
clusion in vol. (S of the continuation of
Peters' Check-list of the Birds of the World
(here usually referred to as "Peters"). The
last classification of the whole family was
that of Hellmayr (1927), and in the inter-
vening 50 years much that requires assimi-
lation has been learned about the family.
David Snow of the British Museum ( N.H. )
has made similar studies of the Cotingidae
and Pipridae, and his results have been
published in two recent papers (Snow 1973,
1975).

Before his death in 1957, John T. Zimmer
almost completed a manuscript of the Ty-
rannidae for Peters' Check-list. However,
Zimmer was concerned almost entirely with
species and subspecies, and, with only
minor exceptions explained in his earlier
published "Studies of Peruvian Birds"
( 1936-1941 ) , he followed the classification
of Hellmayr. While Zimmer's manuscript
is invaluable for defining the limits of spe-
cies and subspecies and their geographical
ranges, he had little to say about the genera
and higher categories. The present paper
deals with those higher categories, and
presents the rationale for the classification
that will be followed in Peters. A subse-
quent paper will consider problems at the
species level.

Bull. Mus. Comp. Zool., 148(4): 129-184, November,

19'

129

130 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

No study of a major family such as the
Tyrannidae would be possible without the
active assistance and cooperation of one's
colleagues. Ernst Mayr, one of the editors
of Peters' Check-list, first suggested that I
edit x'olume 8, which includes the Tyran-
nidae, and he has been a source of constant
support and encouragement throughout. I
have also been fortunate in having Emmet
R. Blake as a colleague in residence at
Field Museum, with whom I could discuss
the innumerable knotty points that arose.
In order to examine the genera and species
not available in Chicago, I have visited
several museums and I would like to thank
for their unfailing courtesy: Wesley Lanyon
and Lester Short of the American Museum
of Natural History, Raymond A. Paynter,
Jr. of the Museum of Comparative Zoology,
and Frank Gill and James Bond of the
Academy of Natural Sciences, Philadelphia.
Kenneth Parkes of the Carnegie Museum,
Pittsburgh, and George Lowery and John
O'Neill of the Museum of Zoology, Louisi-
ana State University, generously lent me
necessary material for examination. I was
fortimate in having David Snow accept
responsibility for the Cotingidae and Pip-
ridae, for we were able to reach complete
accord on the limits of our respective
families. Fran9ois Vuilleumier, John Fitz-
patrick and John Weske, in both corre-
spondence and personal coversations, have
given me the benefit of their field experi-
ence with many of the flycatchers, par-
ticularly the Andean forms, and W. John
Smith has not only done the same, but has
given me a copy of his notes on his own fly-
catcher study. John Fitzpatrick allowed me
to see a copy of his manuscript on the genus
Todirostnim and allies before publication,
and has given me pemiission to reprint one
of the figures from the published (1976)
version. I have benefited greatly from con-
versations and correspondence with Allen
Phillips, primarily concerning the species
of the genus Tyramms. And finally, I am
especially grateful to Stuart Warter for per-
mission to freely use his Ph.D. thesis on

the classification of the Tyrannoidea — one
of the few papers actually defining the
Tyrannid subfamilies — and to publish, as
Appendix D, some of his figures and tables.
Eugene Eisenmann and John Fitzpatrick
were kind enough to critically read the en-
tire manuscript, and to make numerous
suggestions for both fonn and content. I
deeply appreciate their efforts to make this
a more literate and comprehensive publica-
tion. David Snow also read it, to make sure
our concepts of the Cotingidae and Tyran-
nidae were reconciled. I also owe thanks
to my wife Marjorie who read the manu-
script from the point of view of the intel-
ligent layman and made many helpful sug-
gestions, and to Dianne Maurer who typed
it more times than we care to remember.

MATERIALS AND METHODS

External Morphology. During this study
I attempted to use evidence from every
available source^ — external morphology,
anatomy, behavior and distribution. I per-
sonally examined and measured specimens
of every genus recognized by Hellmayr, and
of those described since the publication of
his list. For every genus and for 279 out of
374 ± 1 recognized species, I have plotted
the ratios tail/wing, culmen/wing, tarsus/
wing and culmen ^tarsus as functions of
wing length. I selected these ratios as those
best suited for comparing the proportions of
different sized birds. While wing length is
by no means an exact measure of absolute
size, it seems the most reliable of the
standard measurements.

The different states of various other char-
acters were tabulated for all species. These
were: the presence or absence of abnormal
primaries, and, if present, in which sex;
presence or absence of wing-bars and some
measure of their contrast; presence or ab-
sence of a dark patch at the base of the
secondaries; wing shape, measured by not-
ing the inner primaries to which the 10th
and 9th are equal in length, and by calcu-

Classification of Tyrant Flycatchers ♦ Trayior 13]

lating the ratio of the wmg tip (longest
minus shortest primary) to the wing length;
type of tarsal scutellation; type of nest;
weight by sex; and presence or absence of a
bright crest.

Besides the individual characters listed
above, the degree of overall resemblance
based on plumage color and pattern,
general form and size, shape of bill, extent
of rictal bristles, etc., is still important in
any classification. Such resemblances and
differences have been implicit in previous
classifications and revisions, even though
often subordinated to more artificial key
characters. However, convergence in plum-
age and form is a constant problem, par-
ticularly among the smaller, forest-living
species, and general resemblance alone is
unreliable for diagnoses of genera in this
family.

Distribution. The geographic and alti-
tudinal distributions of each genus were
plotted on outline maps and tabulated
according to the following ecological and
geographical divisions:

I. Lowland humid forest

a) Amazonia, including Venezuela

and the Guianas

b) southeast Brazil

c) Colombian-Pacific: tlie humid

forests of Central America,
western Colombia and north-
western Ecuador

II. Other lowland tropical

a ) woodland/campo/savanna,

south of Amazonia

b) woodland/llanos, north of Ama-

zonia

c) arid: Caribbean coast, Pacific

coast of Peru, etc.

III. Montane forest

IV. Temperate savanna/puna/paramo

a ) montane

b) lowland Chile and Argentina

V. Specialized: lacustrine, marshes,
streams, etc.

VI. Central America, including tropical
Mexico

VII. North America, including temper-
ate Mexico

Literature. Considering that fifty years
have passed since the publication of Hell-
mayr's list, the literature relevant to the
higher classification of the Tyrannidae is
surprisingly sparse. Zimmer ( 1936-1941 )
discussed in great detail the species of fly-
catchers occurring in Peru, which include a
surprisingly large percentage of those of
South America, but his concern was almost
entirely with species and subspecies. In his
1955 manuscript, he followed Hellmayr's
sequence, although he dropped tlie sub-
familial classification. Zimmer's unpub-
lished list was the basis of Meyer de
Schauensee's treatment of the family in his
The species of birds of South America
( 1966). In his A guide to the birds of South
America (1970), Meyer de Schauensee
transferred to the Tyrannidae a few genera
formerly in the Cotingidae, but otherwise
made no changes.

Ames (1971), in his study of the syrinx
of passerine birds, examined 128 species in
86 genera of Tyrannidae. He worked within
the framework of Hellmayr's classification,
that is, he chiefly compared each genus
with others in the same subfamily. In his
taxonomic conclusions, Ames outlined
seven groups of genera (Appendix C)
whose members were more closely related
to each other than to any genus outside the
group, and in some cases he was able to
indicate the degree of relationship between
the groups. However, he did not attempt to
place the groups in a taxonomic hierarchy,
and he was left with a residue of thirty-one
genera that were neither part of nor allied
to any group.

The single paper that directly concerns

132 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

itself \\'itli the elassiiieation ot the Tyran-
nidae as a whole is Stuart L. Warter's
(1965) unpublished Ph.D. thesis on the
cranial osteology of the Tyrannoidea. War-
ter examined the skulls of 160 species in 84
genera of the flycatchers. He concluded
that cranial characters alone did not sup-
port the subdivision of the Tyrannidae into
subfamilies, but he combined these char-
acters witli the external morphological
classification of Hellmayr to produce a
tentative classification that differed in
several significant ways from Hellmayr's.

A number of authors have dealt with one
or the other of Hellmayr's subfamilies, or
parts thereof. Meise (1949) reviewed the
subfamily Tyranninae, and separated them
into three groups of genera primarily on
the type of nest. W. John Smith ( 1966) re-
viewed the genus Tyrannus (including
Muscivora) primarily from a behavioral
point of view, and in 1971 did the same for
Hellmayr's Serpophaginae. Smith and
Francois Vuilleumier ( 1971 ) reviewed the
ground-tyrants, Ochthoeca, Xolmis (includ-
ing Mijioilieretes, Cnemarchiis and Och-
thodiaeta), Neoxolmis, Agriornis and Mus-
cisaxicola; in their study Smith empha-
sized the behavioral characters while
Vuilleumier used the morphological and
distributional evidence. Most recently, John
Fitzpatrick (1976) has reviewed that part
of Hellmayr's Euscarthminae including
Todirostnim and its close allies.

Information on behavior and nest types is
scattered through the literature, the one
exception being Alexander Skutch's (1960)
life histories of Central American birds.
Records of weights are even more scattered,
although a fair amount of data was gleaned
from specimen labels.

The value of the various lines of evidence
cited above proved extremely variable
when applied to the higher classification of
the flycatchers. No single character or
group of characters was consistent through-
out the family. Proportion.s — particularly
the relative length of the tarsus, which was
one of the original characters used in de-

fining subfamilies — vary so much within
Hellmayr's subfamilies that they are with-
out value above the generic or generic
group level. Such characters as abnormal
primaries, bright coronal crests, wing
shape, and wing-bars may be regular in
some genera or groups of genera, but ap-
pear sporadically and irregularly in others.

The type of tarsal scutellation, which
was used by both Sclater (ISSS: 2) and
Ridgway (1907: 328 ff.) as a key family
character, proves so variable that by itself
it cannot even be used to define genera.
Only when this scutellation correlates well
with other characters can it be said to have
taxonomic value. Plotnick and Pergolani
de Costa ( 1955 ) examined the tarsi of some
12,500 passeriform specimens, and pub-
lished figures showing that the classical
exaspidean, pycnaspidean, taxaspidean and
holaspidean tarsal types are not discrete
entities, but blend into each other through
various intermediate forms.

The importance of the syrinx as a taxo-
nomic character is most difficult to assess.
According to Ames ( 1971 : 158 ) , the groups
of genera he defines show a high degree of
syringeal homogeneity and each possesses
certain featiu-es not found elsewhere in the
family. I believe the homogeneity shown
within these groups is real and a true indi-
cator of relationship, because these genera
groups correlate well both with the tradi-
tional classification and with certain inno-
vations suggested by Warter ( 1965 ) on the
basis of cranial characters. On the other
hand, some genera, which on almost every
other character belong in a given group,
have syringes that are imlike those of their
apparent relatives. Annulinicola is not in-
cluded in the Fluvicola group by Ames,
even though the two genera are so close
that I merge them; Todirostnim stands by
itself, although Idioptilon, with which
Short ( 1975 ) merges it, is designated by
Ames as a member of the Colopteryx
group; and Suhlegatus, which is distinguish-
able from Elaenia only by the swollen bill
and lack of a white crest, stands completely

Classification of Tyrant Flycatchers • Trmjlor 133

apart on syrinx from the Elacnia group.
While close resemblance in the syrinx may
be accepted as indicating phylogenetic re-
lationship, the lack of such resemblance
apparently does not negate relationship.

The cranial characters used by \\^arter,
the forms of the nasal and interorbital
septa, and of the palatines and the cranium,
seem to show the greatest consistency in
defining taxa above the generic level.
These characters not only correlate with
each other but \vlth many other characters
as well. In only a few cases does the pre-
ponderance of evidence from other sources
cause me to go against the evidence of the
cranial characters. However, the different
cranial characters vary in importance from
group to group, and no one of them is con-
sistent throughout the family. For example,
among the Fluvicolinae and Tyranninae
the form of the nasal septum is almost 100
per cent consistent, but among the smaller
fl> catchers it is quite variable, and the form
of the orbital septum is diagnostic.

Methods. Because of the variabihty
shown by all the available criteria, I have
not been able to quantify their \alues in any
consistent way. In any given situation, I at-
tempted to use those characters showing the
strongest correlations and to ignore single
contradictory characters, even though in a
different situation the latter might be diag-
nostic. In other words, my approach has
been intuitive, even though I believe it has
also been objective. \Miether the resulting
classification is a valid one can only be
determined by time and by many more
detailed anatomical studies of the whole
family.

In the following discussion I used, for
the sake of simplicity, the generic names
accepted by Meyer de Schauensee ( 1966 ) ,
even though the authors cited ha\'e used
different ones. I used Meyer de Schauen-
see's work in preference to Hellmayr's, be-
cause the former's names are now in
general use and are more familiar to the
majority of ornithologists. Appendix B

lists all generic names whose status has
changed at all since Ilellmayr (1927) and
names that have been proposed since that
date. Use of the appendix sh(juld resolve
most confusion.

LIMITS OF THE FAMILY

In the most recent widely-used classifica-
tion of the perching birds, Passeriformes
(Wetmore, 1960), the Tyrannidae are a
family within the super-family Tyran-
noidea, suborder Tyranni, order Passeri-
formes. They share their superfamily with
the New ^^'orld Cotingidae, Pipridae, Oxy-
runcidae, and Phytotomidae, and with the
Old World Pittidae, Acanthisittidae and
Philepittidae. This is also the classification
of Mayr and Amadou (1951), except the
latter authors use the name "Passeres" for
the order, and make the Oxyruncidae a
sul^f amilv of the Tvrannidae. Ames ( 1971 :
153), on the evidence of the syrinx, re-
moved the three Old \Vorld families to a
position "sedis incertae," and placed the
Tyrannidae, Cotingidae, Pipridae, Oxyrun-
cidae and Phytotomidae in their own sub-
order Tyranni. Despite the shifts in higher
classification the limits of the latter five
families have been remarkably persistent,
remaining essentially the same since the
mid-nineteenth century.

Mayr and Amadou merged Oxyriinctis in
the Tyrannidae in a parenthetical aside,
and did not discuss any characters. Ames
( 1971 : 163 ) treated the Oxyruncidae as a
separate family. He found that Oxyiuncus
had a typically t>aannid syrinx, but no
close resemblance to any particular tyran-
nid genus. The musculature was similar to
that of Pachyrcnnphus in the Cotingidae,
but the cartilages showed substantial dif-
ferences. Waiter was even more positixe in
keeping Oxynincus out of the Tyrannidae.
He stated, "The uniqueness of the highly
specialized [Oxyrunciis] skull argues
against the inclusion of the genus in an
otherwise so relatively homogeneous a
family as the Tyrannidae." I shall follow

134 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

Ames and Waiter in keeping O.xyriincus in
a separate family.

Althougli the families of New World
Tyrannoidea have remained essentially the
same for the past centnry, there has been
some transference of genera and species be-
tween them. In 1907 Ridgway (p. 339)
transferred several taxa from the Tyran-
nidae to the Cotingidae and Formicariidae
because they had types of tarsal scutella-
tion other than the exaspidcan that he con-
sidered characteristic of the Tyrannidae.
These genera were Stigiuattira, Hapalo-
cerctis (= Eu.scorthmus), Hahrnra (= FoUj-
stictiis), Miisci<i,raUa, Culicivora, Sijristes,
Jdiotricctis (= Acrochor(l()))us). Ehiinopsis
(= Mijiopa<iis <i,(iinuir(}ii), Tyrannulus, Mi-
crotriccus and Hylonax ( = Mijiarchus
validiis). However, Hellmayr recognized,
as did Ridgway (1907: 336, ftn.) in an-
other context, that the forms of the tarsal
envelope were of dubious value beyond the
classification of genera, and sometimes not
even there, and he returned (1927) all
these genera to the Tyrannidae. Both War-
ter (1965: 37) and Ames (1971: 162) sup-
ported Ilellinayr's conclusions.

More recently Ames (1971) and Snow
( 1973) recommended transferring from the
Cotingidae, where they have traditionally
resided, to the Tyrannidae the genera
Attila, Pscudattihi, Casiornis, Loniocera
and Rhijtiptcrna. Ames believed syringeal
characters linked these genera into a closely
related group, of which the flycatcher
genus Myiarchus was an integral part.
Snow concurred that the five genera did
not belong in the Cotingids, although he
recognized that so little was known of their
behavior, except for AttiJa, it was difficult
to reach any conclusions. Warter (1965:
37) also placed Rliytipterna in a natural
group with Myiarchus. However, he con-
sidered Attila, while definitely not a Cotin-
gid, sufficiently distinct from the other fly-
catchers as to fonn a subfamily of its own
within the Tyrannidae. Warter did not ex-
amine Casiornis or Laniocera, but he
assumed the former was allied to Attihi,

and the latter to Rhytipterna. These five
genera, based on the then unpublished
work of Ames and Warter, were included in
the Tyrannidae by Meyer de Schauensee
(1970) in his Guide to the Birds of South
America, and by Wetmore (1972) in his
Birds of the Republic of Panama; in both
cases they were placed next to Myiarchus.

Wetmore (1972: 446) also included the
Cotingid genus Lipaugus in the flycatchers.
The genera Laniocera, Rhytipterna and
Lipaugus show a remarkable parallelism in
the geographical distribution of plumage
types. Each genus has a gray species in-
habiting Amazonia, and a representative
rufous species in Central America and
western Colombia. The three genera have
always appeared together in linear lists,
except for that of Meyer de Schauensee
(1970). Snow (1973: 8) remarked on this
parallelism, but felt that on behavioral and
other plumage characters, Lipaugus should
be retained in the Cotingas. Ames lacked
adequate material of this genus. Warter
(1965: 137) found the skulls of Lipaugus
typically Cotingid, and placed the genus in
the subfamily Querulinae. I include Attila,
PseudattUa, Casiornis, Laniocera and Rhy-
tipterna in the Tyrannidae, but leave
Lipaugus in the Cotingidae.

The final genus added to the Tyrannidae
since Hellmayr (1927) is Corythopis.
Corythopis has been considered to form
with Conopophaga a separate family, Cono-
pophagidae, allied to the Formicariidae
and Rhinocryptidae, and included with
them in volume 7 of Peters' Check-list
(1951). Ames, Heimerdinger and Warter
(1968) introduced evidence from the ster-
num, the syrinx, the pterylosis and the
antorbital osteology showing Corythopis
belongs in the Tyrannoidea — not with
Conopophaga, or with the Formicariids,
with which the authors unite Conopophaga.
Within the superfamily, the syrinx, pterylo-
sis and external appearance are far more
typical of the Tyrannidae than of any other
family. The authors did not determine the
precise relationships of Corythopis within

Classification of Tyrant Flycatchers • Tr&ylor

the Tyrannids, but the pterylosis resembles
that of Helhnayr's Eiiscarthmines. Meyer
de Schauensee (1970: 326) included Corij-
thopis in the Tyrannidae, but placed it at
the end of the family because he was un-
certain of its position. Ames (1971: 67)
inserted Conjthopis between Euscarthmus
and Pseiidocolopteryx, but he described the
syrinx as "unlike any of the above," mean-
ing the subfamily Euscarthminae.

A difficult genus and species to place is
Xenopsaris aUnnucha, which has been
transferred back and forth between the
Cotingids and Tyrannids several times.
The species aUyinucha was originally de-
scribed in the Cotingid genus Fachij-
rinnphus (Burmeister, 1869), and was rec-
ognized as a Cotingid by Sclater (1893),
after he had unaccountably omitted it from
his Catalogue of Birds ( 1888). In the mean-
time, aJhimicha became the type of two
new genera, Xenopsaris Ridgway, 1891,
which the author considered Cotingid and
near the genus Casiornis, and Prospoeitus
Cabanis, 1892, which that author considered
Tyrannid and near the genus Serpopha^a.
Berlepsch (1907) and Hellmayr (1927)
kept Xenopsaris in the Tyrannidae, but
Zimmer, in his Peters manuscript, removed
it to the Cotingids as a subgenus of Pachij-
ranipJuis. Meyer de Schauensee (1966)
placed Xenopsaris next to PacJiyranipJuis
but as a separate genus. Finally Smith
(1971) tentatively placed it again in the
Tyrannidae, in the subfamily Serpophagi-
nae, as did Snow ( 1973 ) . Unfortunately,
neither Ames nor Warter had specimens.

The characters of Xenopsaris that most
suggest relationship to the Cotingids are
its taxaspidean tarsus and the glossy black
crown in the male, which is like that of
many PachyranipJius. Parkes (in lift.) con-
siders that its slightly swollen bill with pale
cutting edge suggests Pachyramphus, and
that its sexual dimorphism is similar to that
of Pacliyraniplms but has no counterpart
among the Serpophagine flycatchers. On
the side of Tyrannid relationship, Xenop-
saris males lack the shortened and pointed

9th primary that is characteristic of males
of all Pachyraniphus species; in size Xenop-
saris is smaller than any Pachyraniphus,
much nearer the size of the Serpophagines;
it builds a cup-shaped nest rather than a
large globular structure with side entiance
characteristic of the Becards; its vocaliza-
tions are unlike those of Pachyraniphus;
and the white outer edge of its outermost
rectrix is a common condition in Tyrannids,
but not found in Pachyraniphus. Xenop-
saris' palustrine habitat would be unique
among the Cotingids, but obviously not im-
possible.

I believe the differences in the 9th pri-
maiy and in the nest form remove Xenop-
saris from any close relationship to Pachy-
raniphus, and that plumage similarities are,
therefore, the result of convergence. This
leaves only the taxaspidean tarsus and the
bill form and color noted by Parkes (in Utt.)
as characters linking Xenopsaris with the
Cotingids. \Miile taxaspidean tarsi are
characteristic of the Cotingids, they are also
found in some Tyrannid genera such as
Culicivora, Stigniatura and Inezia. The bill
of Xenopsaris is more slender than that of
Pachyraniphus, which is the reason Ridg-
way and Cabanis placed it near Casiornis
and Serpophaga respectively, and the color
of the bill is much like that of some species
of the Tyrannid genus Knipolegus, al-
though unlike any of the Serpophagines. I
consider Xenopsaris a Tyrannid, but be-
cause of the differences in sexual dimoiph-
ism and bill color I believe the resemblance
to the Serpophagines, particularly to Serpo-
phaga cinerea, is due to convergence. Since
Xenopsaris is of uncertain relationship
within the Tyrannids, I shall place it at the
end of the family incertae sedis.

Warter (1965: 97-100; 13.8-140) dis-
cusses the cranial characters of the Cotin-
gid Tityras [Tityra and Erator, the latter a
genus not recognized by Meyer de Schau-
ensee (1966: 320)] and Becards (Pachyrani-
phus and Phitypsaris). Both groups have
essentially Tyranno-Myiarchine skulls, but
differ from an>' of the recognized Tyran-

136 BiiUctin Museum of Comparative Zoology, Vol. 148, No. 4

nids in a number of characters associated The Tityrinae, however, are tentatively
with the nasal capsule. The two Tityras, allied to the Tyrannidae only because their
caijmia and semifa.sciata, are the most crania more nearly resemble those of the
al:)errant. According to Warter (p. 99), Tyrannids than those of the Cotingids. In
"The peculiar structure of the tityrine bill Peters' Check-list, the Tityrinae will be
and nasal capsule . . . provides a character placed at the end of the Tyrannidae, and
which, by its veiy uniqueness, constitutes they will not be considered further in the
a radical departure from an essentially discussion of subfamilies below,
conservative pattern that obtains through- Two genera of Pipridae have recently
out the tyrannoid series." The skull of Era- been suggested as possible members of the
tor inquisitor, however, despite the very Tyrannidae. Warter (1965: 133) felt that
close resemblance of that species to Tityra Neopelma should properly be in the Tyran-
in plumage, external morphology, voice and nids, and Ames (1971: 160) recommended
nesting habits, is not so extreme, and shows placing Piprites with his Myiobius group of
more resemblance to the Becards. The lat- Tyrannids. Warter considered Neopelma
ter, in turn, seem more closely related to one of several Piprid genera that seemed
the Tyrannids. Ames (1971: 163) also intermediate between Pipridae and Tyran-
found that the syringes of the Becards had nidae, and the one most closely resembling
several Tyrannid features not found in the the Tyrannids. He did not, however, ally it
Cotingas. to any given genus or subfamily. Ames did
Warter (pp. 139-140) suggested a num- ally Piprites to his Myiobius group, includ-
ber of alternatives for treating the Tityras ing Myiobius, Terenotriccus, Pyrrhomyias
and Becards. The first possibility was to and Onychorhynchus; this syringeal type
emphasize their distinctiveness from the was so distinctive that Ames (p. 122)
remainder of tlie Cotingas by recognizing placed his Myiobius group in a different
them as a family Tityridae. If further study structural division from the remainder of
should show a closer link ])etween the the Tyrannidae, along with Piprites and the
Becards and the Tyrannidae, the fomier majority of the Cotingas. However, Pip-
could be made a subfamily, Pachyram- rites, with its short, deep bill, stout build,
phinae of the Tyrannidae, and the separate short tail and sexual dimorphism, is so un-
iamily Tityridae maintained for Tityra and like any members of his Myiobius group,
Erator. However, he thought the "most that I think the syrinx resemblance is due
logical" action would be to ally the subfam- to convergence. Unfortunately Ames did
ily Tityrinae to the Tyrannidae, recogniz- not have a specimen of Neopelma, nor did
ing within it two tribes, Tityrini and Pachy- \\'arter have a skull of Piprites, so it is not
ramphini. Although I accept his conclusions possible to determine if there is a coiTcla-
that the Tityrinae may be closer to the fly- tion between the syringeal and cranial
catchers than to the Cotingas, I do not characters. Snow (1975: 22) recognized
consider them the hierarchical equivalent that Piprites might be related to the Tyran-
of the other Tyrannid subfamilies that I do nids, but he preferred to keep it in the pip-
recognize, the Elaeniinae, Fluvicolinae and rids for the time being. In view of the
Tyranninae. The skull uniformity among uncertainty in both cases, I shall leave Neo-
the Tyrannidae of Hellmayr and previous pelma and Piprites in the Pipridae, where
authors does not support the recognition of they are both readily retrievable,
subfamilies on cranial characters alone
(Warter 1965: 131), and the Elaeniinae, CLASSIFICATION TO SUBFAMILIES

Fluvicolinae and Tyranninae are based on a Hellmayr ( 1927) divided the Tyrannidae

combination of cranial, syringeal, external into seven sul)families: Fluvicohnae, Tyran-

ni()ri")li()logical and behavioral characters, ninae, Myiarchinae, Platyrinchinae, Eus-

Classificatiox of Tyrant Flycatchers

.rai

caithminae, Serpophaginae and Elaeniinae.
Hellmayr did not define his taxa, and be-
fore they are critically examined, some
attempt must be made to determine what
criteria he had in mind. This is not an easy
task, nor possibly even a profitable one. As
Warter (1965: 130) remarked when dis-
cussing family group taxa, "Either the
originality of, or the precedent for, the taxa
used is often impossible to ascertain." How-
ever, a brief history of the development of
Hellmayr's subfamilies since Sclater ( 1(S88 )
— the last author to review the whole family
and to characterize his subfamilies — may
give us some idea of what the former had in
mind.

Fluvicolinae - This is the one subfamily
that has descended with comparatively
little change from Sclater. He characterized
it as "Feet strong, tarsi stout, elongated;
habits more or less terrestrial; coloratic^n
gray, white and black." Morphologically it
is extremely heterogeneous, but the species
agree in being non-forest types with their
center of distribution south of the Amazon
and in the temperate zones of the Andes,
Argentina and Chile. Ihering (1904: 320)
suggested removing Sayornis because of its
North American distribution, but this was
ignored by Berlepsch (1907) and Hellmayr.

Tyraxxixae - Sclater characterized this
subfamilv as "Feet strong; tarsi short but
strong; habits arboreal but frequenters of
more open spaces; coloration olive, gray,
white or yellow." Although the name per-
sists, the composition of the family has
changed drastically. It originalh' contained
only four of Hellmayr's Tyranninae genera,
the remainder of which were in the Elae-
niinae, but it also had all the genera of
Hellmayr's Myiarchinae. Ihering (1904:
318) recognized a subfamily Pitanginae
(Hellmayr's Tyrannine genera Legatus
through Pitongiis), which he moved from
the Elaeniinae to the vicinity of the Tyran-
ninae, with which they were incorporated
by Berlepsch (1907: 473) and Hellmayr
(1927). Ihering's reasons for the shift were
the wide distribution and aggressive be-

havior of the Pitanginae. Hellmayr's Tyran-
ninae took final shape when Berlepsch
remo\-ed those genera now in the Myiarch-
inae.

Myiarchixae - Set up as a new subfamily
by Berlepsch (1907: 476), but nowhere
characterized; composed of genera formerly
placed by Sclater in the Tyranninae.

Platyrixchixae - Characterized by Scla-
ter as "Feet weak, tarsi thin; forest dwel-
lers; coloration olive and yellow; bill
depressed; rictus bristled." Of the 20
genera included by Sclater, only the
nominate genus appears in Hellmayr's sub-
family! Ihering (1904: 318, 321) dismem-
bered Sclater's family, placing those genera
that made a pendent, pyriform nest in a
new subfamih', Euscarthminae, and those
making a cup-shaped nest in another new
subfamily, Seqoophaginae. Berlepsch (1907:
482) recognized Ihering's Euscarthminae,
although calling it Platyrinchinae, but ex-
tracted from it three genera for which he
created a new subfamily, Rhynchocyclinae.
It was the latter family, to which was
added Plati/riitchus\ that became Hell-
ma>'r's Plat\'rinchinae.

Euscarthmixae - As noted above, this
family was first recognized by Ihering as
those genera of Platyrinchinae that made
pendent, pyriform nests and were primarily
forest forms. It was called Platyrinchinae
by Berlepsch and then Euscarthminae
again by Hellmayr when he removed Platy-
rinchus. Unfortunately, the name Euscarth-
minae was there incorrect, because Ihering's
type genus Euscarthnuis was called Eus-
carthmorms by Hellmayr, and he used the
name Euscortliinus for the taxon known as
Ilapalocerciis by Ihering, a cup-nest
builder!

Serpophagixae - A family created by
Ihering for the genera of Sclater's Platy-
rinchinae that built open, cup-shaped nests
and inhabited open coimtry or the Andes.
It was accepted b>^ Berlepsch (he actually
called it siibfani. nov.) and by Hellmayr.

Elaenuxae - The last of Sclater's sub-
families, which he characterized as, "Feet

138 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

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Classification of Tyrant Flycatchers

1 fC'j

'ii ■^-> '

weak; tarsi thin; forest dwellers; coloration
olive and yellow; bill compressed; rictus
smooth." After Ihering (1904: 318) re-
moved the Pitanginae — "large birds with
strong bills that biologically much approxi-
mate to true Tyranninae" — and Rhyncho-
cijclus, because of its pendent nest, the
subfamily was accepted unchanged by Ber-
lepscli and Hellmayr.

These changes are summarized in Table
I.

The families of Sclater, except for the
Fluvicolinae, h\\\e lieen so stronglv revised
that his original characters are no longer
pertinent. Even Sclater's characters for the
Fluvicolinae are irrelevant, for such genera
as Muscipipra have among the shortest tarsi
in the family instead of "stout, elongated."
The Myiarchinae were never defined, and
it is hard to know what Berlepsch had in
mind, because they vary from some of the
larger Myiarchus, with wings 90-100 mm to
Terenotriccus with wing 52 mm, and from
the small, round winged, forest haunting
Myiubius, to the large, open-country,
pointed winged Hiniiiditiea. Ihering's sep-
aration of the Euscarthminae from the
Seipophaginae on the basis of nest type
was a major advance, because this char-
acter correlated well with proportions of
I bill and tarsi, but there were, and still are,
^ so many problem genera for which the nest
i type is unknown, that its usefulness is
limited.

Certainly mensiu'al characters do not in
any way define the present subfamilies.
Taking wing length as a criterion of size,
all the Tyranninae are larger than any of
the Euscarthminae and Serpophaginae;
otherwise there is overlap among all the
subfamilies. The same is true of propor-
tions. A species with a wing 62 mm, tail
52 mm, culmen 14 mm and tarsus IS mm
would fit in any subfamily except the
Tyranninae without distorting the present
limits of the taxon. It is clear that the
present subfamilies are based on such qual-
itative characters as the form of the bill,
the extent of the rictal bristles, the shape of

the wing and tail, habitat, distribution,
color and pattern. This does not imply that
such a classification is valueless; Sclater,
Ihering, Berlepsch and Hellmayr were all
highly capable and experienced ornitholo-
gists, and the classification they developed
improved at each stage. However, it is
futile to search their work for key mensural
or moiphological characters that would
clearly define their subfamilies.

In the literature since Hellmayr, there
have been no attempts to evaluate his sub-
families. Ames ( 1971 ) used the subfamilies
of Hellmayr as the framework within which
he conducted his investigation of the syrinx.
His genus-to-genus comparisons were al-
most all intra-subfamily, and although he
set up groups of related genera, he ex-
pressly avoided equating them with sub-
families. Zimmer ( 1936-1941 ) in his
Studies of Peruvian Birds, and also in his
preliminary manuscript for Peters, ignored
subfamilies. He could hardly have studied
the Peruvian flycatchers, which include
over half the known species, without form-
ing some ideas about subfamilies. How-
ever, no hint of them appears in his works,
and he followed the order of genera in
Hellmayr's list without change. In one
instance one gets the impression that he
doubted the reality of some of the subfam-
ilies completely. When he (1940b: 19)
transferred Mecocerculus superciliosus and
Leptopogon nigrifrons to the genus Phyllos-
cartes, he merely stated "there are no posi-
tivelv distinguishing characters between
these two species and various .species of
PJiylJoscartes" and he did not mention the
fact that the genera had been assigned by
Hellmayr to three different subfamilies,
Serpophaginae, Elaeniinae and Euscarth-
minae respectively.

The one study that discusses the Tyran-
nidae at the subfamily level is that of Wai-
ter (1965). Although his cranial characters
alone did not support the separation of the
Tyrannidae into subfamilies they did,
when combined with the classification of
Hellmavr based on biological and external

140 Bulletin Museum of Coinporativc Zoology, Vol. 148, No. 4

morphological characters, suggest a revised
classification that Warter (p. 131) pre-
sented "merely as a model for other
workers."'

In his study of the Tyrannoidea, Warter
used five cranial characters:

a) nasal septum - six forms of the nasal
septinn were recognized (only five occurred
in the Tyrannidae), which were determined
by the type of supporting structure and by
the presence and form of the transverse
plate.

b ) interorbital septum - five states of the
interorbital septum were recognized (only
four in the Tyrannids), characterized by
the number and extent of the fenestrae.

c) prepalatines - three states, deter-
mined by their flattening and curvature.

d ) cranium - three forms of the cranium
depending on overall configuration.

e) palatomaxillaries - presence or ab-
sense recorded.

In his Figures 2 and 3 (Appendix D),
Warter illustrates the states of his cranial
characters.

Warter examined the crania of S50 speci-
mens of flycatcher, representing 160 spe-
cies in 84 genera. In his Table II (Appen-
dix D) he lists the state of each of his five
characters by genus. The Tyrannidae as a
whole are homogeneous and there were too
few long series to adequately investigate
individual variation at species level; there-
fore his treatment had to be at the genus or
group of genera level (p. 93). This prob-
lem did not exist among the more hetero-
geneous Cotingas, where single skulls of
the genera and even some of the species,
may be readily identified. In his Table II,
Warter lists the genera in the order of llell-
mayr.

Warter's Table II as reprinted here
(Appendix D) differs slightly from the
form in which it appeared in his disserta-
tion. With his permission, I have included
several ambiguous entries from an earlier
draft that were eliminated from the final
copy. These additions have been enclosed

in parentheses; figures outside the paren-
theses, including the question marks, are
those of his final version. The ambiguities
are consequent upcni the difficulty of assign-
ing characters to a given type because of 1)
shot damage, 2) careless cleaning, 3) overly
zealous dermestids or 4) a continuum of
form requiring arbitrary assignment to one
type or the other. These ambiguous states
are included here to help evaluate which
data are equivocal, and which relatively
certain.

When Warter's characters are compared
with Ilellmayr's classification, there is a
strong correlation between the individual
character states, and sometimes complexes
of states, and the broad groupings of
genera. This inspires confidence that both
the evolved classification of Hellmayr and
Warter's cranial characters reflect real re-
lationships among the Tyrannidae, and not
just superficial resemblance or convergence.
The most striking example is found in the
Tyranninae, all species of which have a
character complex including type 1 or 2
nasal septum without basal plate, and types
1 interorbital septum, palatines and cra-
nium. It is highly improbable that such a
complex of four characters would have
evolved independently more than once,
thus the Tyranninae are almost certainly a
monophyletic group. Since the cranial
characters do correlate so well overall with
what is known of Tyrannid relationships,
they should be given considerable weight
when they diverge from the old classifica-
tion.

Warter (p. 94) found that the three sub-
families of "larger' flycatchers, Fluvi-
colinae, Tyranninae and Myiarchinae, lent
themselves better to characterization than
the four remaining subfamilies of "smaller"
flycatchers. Although, as noted before,
there is overlap in size among almost all
subfamilies, the latter four of Hellmayr —
Platyrinchinae, Euscarthminae, Serpophag-
inae, and Elaeniinae — have a preponder-
ance of small species, for which skeletal
material is less common, and, being deli-

Classification of Tyrant Flycatchers • Traylor 141

cate, are frequently damaged, \^^arter (p.
96) stated, "Perhaps largely as a result of
insufficient material, the adequate char-
acterization of the smaller flycatchers, is,
at the present time, impossible."

Within the larger flycatchers, there is a
clear dichotomy. The Tyranninae of Hell-
mayr are a distinct group as outlined above.
Belonging to this group, as shown by
identical cranial characters, are the genera
Mijiorchus and Eribates of Hellmayr's
Myiarchinae, and Rhytipterna, placed by
Hellmayr and his predecessors in the
Cotingidae. Presumably part of the same
complex, although Warter had no material
of them, are Hylonax, which has been syn-
onymized with Myiarchus by Lanyon
(1967b: 339); DeJtarhynchus, which is a
Myiarchus with a short broad bill; and
Laniocera, which has been considered a
close relative of Rhytipterna except by
Ridgway ( 1907 ) , who placed it in the Pip-
ridae while leaving Rhytipterna in the Co-
tingidae. The second group of large fly-
catchers is composed of the Fkuicolinae of
Hellmayr and most of the remaining genera
of that author's Myiarchinae, the genera
NuttaUornis through Myiophobus. This
group is characterized by a type 6 nasal
septum with horizontal baseplate, and
either types 2 or 3 of at least two of the re-
maining three characters; the type 2 inter-
orbital septum occurs almost exclusively in
this group, although types 1 and 3 occur
as well. Warter (p. 95) considers that the
second group's two subgroups, the Fluvi-
colinae and the remainder of tlie Myiarchi-
nae resjjectively, may be distinct on char-
acters too minor to justify separation at the
subfamily level. They merge insensibly
together through such genera as Ochthoeca,
Ochthornis and Entotriccus and may repre-
sent terrestrial or semi-terrestrial and ar-
boreal divisions of the same stock.

A third group that Warter (pp. 95, 140)
feels should be attached to the section of
large flycatchers, either as a subfamily or
"subfamily-equivalent" group are Attila
and the probably closely related Fseudat-

tila and Casiornis, two genera not available
to him. Skulls of Attila were essentially
Tyranno-Myiarchine, but with the follow-
ing differences: interorbital septum less
completely ossified ( type 5, not found else-
where in tlie Tyrannidae); nasal septum
incomplete, shallow, along its entire length;
prepalatines narrow, slightly convergent,
nearly straight. In themselves, the Attila
skulls were a distinct type. Warter's sug-
gestion was to recognize among the large
tyrant flycatchers three "subfamily-equiva-
lent" groups: Attilinae, Tyranninae and
Fluvicolinae.

Warter, as noted before, did not consider
tliat cranial characters permitted the classi-
fication of Hellmayr's four subfamilies of
small flycatchers at this time. He did be-
lieve (p. 131) the Euscarthminae graded
into the Platyrinchinae \da Todirostruni
and Oncostoma, and the Serpophaginae
graded into the Elaeniinae via Serpophaga.
He also included in this section the genus
Onychorynchus, which, like Platyrinchiis,
has a type 3 nasal septum and does not
belong in either section of the Myiarchinae,
the subfamily where Hellmayr placed it.

Warter's (p. 131) final tentative classi-
fication of the Tyrannidae was a com-
promise between the cranial evidence and
the subdivisions of Hellmayr based on bio-
logical and external morphological features.
It included five subfamilies, three of them
divided into tribes:

Attilinae (see W^irter, pp. 95 and 140)

Tyranninae

Fluvicolinae

Fluvicolini

Alectrurini

"Contopini"
Platyrinchinae

Onychorhynchini

Platyrinchini

Euscarthmini
Elaeniinae

Elaeniini

Serpophagini

The classification adopted in this paper

142 Bulletin Mii.sctiDi of Conipurutkc Zoology, Vol. 148, No. 4

is essentially that ot Waiter at the siibhim-
il\' le\'el, except that I do not recognize his
Attilinae, and I consider that all of the
small flycatchers, his Platyrinchinae and
Elaeniinae, belong in a single subfamily for
which Elaeniinae is the oldest name. The
use of tribes, however, does not seem war-
ranted. The three Fluvicoline tribes are un-
even, both in numbers of genera and
desfree of difference, and the Fluvicolini
and "Contopini" may not be so distinct as
previous classifications would suggest. The
tribes of Elaeniinae, which Warter recog-
nized in deference to previous classifica-
tions, seem to merge into each other io
insensibly that no divisions can be recog-
nized.

The brief history of the e\'olution of Hell-
mayr's subfamilies showed that none of
them was maintained on clear-cut charac-
ters, and that the Myiarchinae had never
been diagnosed, not even by the original
describer. The dismemberment of the
Myiarchinae, therefore, may upset our
sense of familiarity with a subfamily of 70
years' standing, but it does not do violence
to any known diagnostic characters. The
more important (}uestion is how well do the
segments of Myiarchinae fit into the Tyran-
ninae and Fluvicolinae respectively.

Myiarchus, with the closely related Eri-
luite.s, Ili/lonax and Deltarhijnchus, fits
easily into the Tyranninae. It is a highly
successful genus, like so many of those in
the Tyranninae, with a geographical range
ec^ual to that of Ti/ranmis itself, from tem-
perate North America to temperate South
America. Form and proportions are typi-
cally "flycatcher," with a long, broad and
moderately flattened bill and about average
development of rictal bristles. The tarsi
are longer than those of most of the Tyran-
ninae, but still short enough that Sclater
put them in with the "tarsus short, stout"
Tyranninae. The hole nesting habits of
MtjUnchus are shared for the most part
with Mijiodtinaslcs. There are no evident
reasons for keeping Mijiarchus out of the
Tyranninae, and the cranial evidence for

placing it there is compelling. The latter
statement is also true for RJiytiptenia and
presumably Laniocerci, and the transfer of
these two genera to the vicinity of Mijiar-
chus on other grounds by Snow ( 1973) and
Ames ( 1971 ) reinforces the cranial evi-
dence. Meyer de Schauensee (1970) and
Wetmore ( 1972 ) followed Ames and War-
ter, whose theses they had seen before
publication.

The recognition of Warter's subfamily
Attilinae, Attila and Casiornis, must be con-
sidered here, because the above authors all
included those genera, previously included
in the Cotingidae, with Rhiitipicrna and
Laniocera next to Myiarchus. Externally,
there is nothing that keeps Attila and Casi-
ornis separate from Rhijtiptcrna, Laniocera
or Myiarchus. Internally, there is strong
evidence from the syrinx that they are
closely related. The above five genera
make up Ames' (1971: 160) Myiarchus
group. Speaking of the former Cotingid
genera compared to MyiarcJms, Ames says,
"Without assuming interdependence of
several syringeal featmes, it is difficult to
see how two groups of birds could evolve
both external and internal similarities to
such a degree." If this is true when Attila,
Casiornis, Laniocera and RJiytipterna are
compared to Myiarchus, it is equally true
when Attila is compared to Rhytipterna
and the others. Despite the fact that Attila
has a partially ossified interorbital septum
not found elsewhere in the Tyrannidae,
which caused Warter to put it in a separate
subfamily, I consider Ames' Myiarchus
group to be monophyletic, and I keep it to-
gether in the Tyranninae.

The transfer of the remainder of the
Myiarchinae, Warter's "Contopini," to the
Fluvicolinae suggests some relationships
not considered before. Both are presum-
ably old lineages with a high degree of
sympatry in the larger genera, numerous
monotypic genera and relict species, and a
high degree of phenetic variability. War-
ter's Fluvicolini contains all the truly terres-
trial flycatchers and is usually thought of

Classification of Tyrant Flycatchers • Traylor i43

as a long-legged group. However, there are
a number of arboreal Fluvicolines, such as
OcIitJioeca and KnipoJeii^us, and these are
very close in proportions to the Contopines.
When the two are treated as related, cer-
tain cases of resemblance between genera
and species that were previously thought of
as convergence may turn out to show close
relationship. A striking example is the
Fluvicoline Ochthoeca diadema and the
Contopine Myiophobus flavicans. Both are
bright olive green flycatchers, identical in
wing length and shape, bill form, and rictal
bristles; they can be separated only by the
slightly longer tarsus and yellow super-
ciliaries of diadema, and the yellow crest of
flavicans. Their resemblance may be due to
the persistance of the characters of an early
common ancestor.

The Fluvicolines of Sclater and Hellmayr
are a heterogeneous assemblage united by
their dry country or temperate center of
distribution in South America. Their great
phyletic age has produced such unlike
forms as the long-legged, long-billed terres-
trial Muscisaxicola and the short-legged,
short-billed arboreal Muscipipra. It is en-
couraging, and a tribute to Sclater's intui-
tion, that Waiters cranial characters sup-
port the present classification. Waiter's
Contopines on the other hand, are a lin-
eage that has had its most successful radia-
tion in North and Central America. While
there was a proliferation of species in the
Fhuicoline genera Muscisaxicola, Aiiriornis
and Xolmis in the south temperate and sub-
tropical regions, there was a similar pro-
liferation in the Contopine Contopus.
Empidonax and Saijornis in the north tem-
perate regions and Middle America. The
Flu\icolines are still virtually all South
American, with only one species reaching
eastern Panama, while only a few genera of
Contopines are primarily South American.
These are mostly small, lowland forms,
such as Myiolyius and Terenotricciis, or
montane forest genera such as Myiotriccus.
Pyrrliomyias and Myiophobus. These South
American genera may have separated from

the North and Central American group
fairly early, because some at least have a
distinctive type of syrinx, as discussed be-
low. However, in general fonn all these
genera are fairly close, being "typical"
small flycatchers with broad, depressed
bills and moderate to heavy rictal bristles.

Three of Ames' genera groups are in-
cluded in the expanded Fluvicolinae. The
first of these, his Fluvicola group, belongs
to what might be called the core of War-
ter's Fluvicolini: Xolmis, Neoxolmis, Af^ri-
oniis, Muscisaxicola, Fluvicola, Gubernetes,
Knipolegus, Muscipipra and Phaeotriccus.
With the possible exception of Gubernetes
and Muscipipra, one would expect these
genera to be alike. What is surprising is
the absence of Arundinicola, Entotriccus
and Yctapa, which are in all morphological
characters except the syrinx very close to
Fluvicola, Phaeotriccus and Gubernetes
respectively. Ames' second group is the
Xuttallornis group — Nutfallornis, Sayornis,
Contopus, Blacicus, Empidonax, AecJimolo-
phus, and Aphanotriccus — which according
to Ames is closest to the Fluvicola group in
syringeal structure. It is this group that has
been so successful in North and Central
America.

The members of Ames' Myiobius group
— Myiobius, Terenotricciis, Pyrrhonujias
and OnychorJiynchus — appear more closely
related to each other than to any other
group, and Ames placed them in a separate
major structural division of the syrinx from
the rest of the flycatchers. This is the group
that is primarily of South American forest
distribution. The inclusion of Onycliorhyn-
cJuis with the Myiobius group agrees with
the traditional classification of Hellmayr,
but differs from that of ^^^u■ter, who con-
sidered the genus an aberrant "small" fly-
catcher in his subfamily Platyrinchinae.
The status of Onychorhynchus is con-
sidered in more detail below.

The remainder of the family, which War-
ter. in deference to previous classifications,
divided into two subfamilies and five tribes,
seems to me incapable of such subdivision.

144 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

Consider first the cranial characters. The
nasal septum, which has proved an im-
portant conservative character in the Fluvi-
colinae and Tyranninae, is extremely vari-
able in the Elaeniinae, and lacks any
correlation with other characters. On the
other hand, his tribes Platyrinchini, Eus-
carthmini and Seipophagini, and some
genera of the Elaeniini, all show a type 4
interorbital septum, which does not occur
at all in the large flycatchers. Of those
Elaeniini genera that lack the type 4 inter-
orbital septum and that might be set apart
on that account, three are linked to the
Seipophagini by their type 5 nasal septum,
which is unique to these two groups. As
Warter said (p. 112) characterization of
the smaller flycatchers on the basis of their
cranial characters is impossible.

The external morphological and bio-
logical evidence for subdividing the small
flycatchers is little better, despite the four
subfamilies usually recognized. Onijcho-
rhynchus, with its striking ornamental
crest, long, flat bill, and long rictal bristles,
is out of place in any group; it fits no better
with the small flycatchers than with the
Myiarchinae, where Hellmayr placed it.
It is seemingly related to Platyrinchus by
the type 3 nasal septum, which is found
only in these two genera, but Platyrinchus
has a short, broad, flattened bill, a modest
concealed crest, and makes a simple cup-
shaped nest rather than the elaborate, pen-
dent, pyriform nest of Onychorliynchus.
Platyrinchus is somewhat out of place in
its own tribe because of its peculiar nasal
septum and simple cup-shaped nest. War-
ter (p. 131) found that on cranial characters
the Platyrinchini and Euscarthmini inter-
graded through Todirostrum and Onco-
sfonui, and that the Seipophagini and
Elaeniini were related through Serpophaga.
On external moiphological and biological
characters I can find no way to character-
ize family-level groups. There are certain
core groups that are (juite distinct, such as
the flat-billed RJiynchocychis type, the

tod\'-flycatchers Todirostrum and allies,
and the small-billed generalized Elaenia
group, but there are more genera falling be-
tween these groups than within them.

Ames' two genera groups from this part
of the family are essentially two of these
core groups. His Colopteryx group — Colop-
teryx, Oncostorna, Idioptilon, Hemitriccus,
Myiornis and Lophotriccus — are all mem-
bers of what I call Todirostrum and allies,
although Todirostrum itself has a distinc-
tive syrinx and is not placed with the
others. His Elaenia group includes Elaenia,
Suiriri, Camptostoma, Tyrannulus and
Phaeomyias, all part of what I call the small-
billed generalized Elaenia group. However,
the Rhynchocyclus type, that I mention as a
core group, is considered by Ames as an
artifact of convergence. Speaking of Platy-
rinchus, Tolmomyias and Rhynchocyclus,
he (p. 161) says that they, "differ so
strongly in their syringeal structure that
one may seriously question the reliability
of bill shape as a common character for the
five genera" [including Cnipodectes and
Ramphotrigon, which he had not seen].
Ames' evidence agrees with that of the
nasal septum and nesting habits to show
that Platyrinchus is out of place even in the
restricted subfamily where Hellmayr put it.

The recognition of any Elaeniinae sub-
divisions, either the earlier subfamilies or
^^^arter"s tribes, would give a sense of cer-
tainty about the reality of these entities that
is non-existent.

The sequence of subfamilies and of
genera in the family lists for Peters' Check-
list ideally go from primitive or general-
ized to advanced or specialized. Histor-
ically, the lineal list of the flycatchers has
begun with the Fluvicolinae, and ended
with the Tyranninae (Sclater, 1888) or the
Elaeniinae (Berlepsch, 1907; Hellmayr
1927). Although there is httle solid evi-
dence, the flycatchers seem most nearly
related to the Pipridae, with Elaeniine fly-
catchers and the sexually monomoiphic
Piprids nearest the ancestral line. Warter

Classification of Tyrant Flycatchers • Traijlor

145

(p. Ill) found the skulls of the Piprids
Tyranneiites and Neopelma flycatcher-like,
and intermediate between the flycatchers
and typical Piprids. This suggested an
intermediate position for the Pipridae be-
tween the Cotingidae and Tyrannidae.
Ihering (1904: 319) came to a similar con-
clusion many years before when he con-
sidered that the Pipridae and Tyrannidae
descended from a common ancestor whose
closest relati\'es were among the Elaeniinae
(sensti stricto) in the Tyrannids and the
Ptilochlorinae (including Tyranncutes and
Neopelma) in the Piprids. The ancestral
fomi was presumably small, oli\'e green,
arboreal and forest-living, with a short,
somewhat compressed and bristled bill; a
description that fits many genera among
the Elaeniinae. Ames (1971: 150) did not
find syringeal characters useful taxonom-
ically in determining relationships among
the higher categories of Tyrannoidea. As he
says (p. 157), "Except for the Piprites-
Myiobius complex and Iliciira, the few
manakins examined show little syringeal
similarit)^ to the topical members of either
the Tyrannidae or Cotingidae."

Assuming that the Elaeniine flycatchers
are the most generalized or primiti\'e, then
the Fluvicolinae, with their manv terres-
trial forais, are the most specialized, and
the Tyranninae are the most "advanced."
I I use "advanced" in the sense of most suc-
; cessful and presumably most recently
; evolved, for not only are the genera widely
spread geographically, from North America
or Mexico to southern South America, but
many individual species are equally wide-
spread. Fitzpatrick ( in litt. ) points out that
wide distributions in the Tyranninae are
almost always associated with open country
and edge habitats, and bear no relation to
recent speciation. However, the extreme
uniformity in cranial characters among the
Tyranninae, and the similarity in external
moiphology among them compared to tlie
diversity in the Fluvicolinae, suggest to me
a more recent proliferation. The sequence

of the subfamilies for Peters' Check-list
will therefore be: Elaeniinae

Fluvicolinae
Tyranninae

SEQUENCE AND RELATIONS OF
GENERA

The sequence of genera that follows
(Appendix A) relies heavily on tradition
when there is no strong evidence to suggest
a change. If I had started de novo, I might
well have made considerable changes.
However, they would have been based on
characters no more conclusive than those of
Berlepsch or Hellmayr. There being no
ad\'antage in producing a new sequence
just for the sake of change, much of the fol-
lowing list follows Hellmayr, but in reverse.

ELAENIINAE

Just as the subfamilies were placed in the
sequence of primitive or generalized to ad-
vanced or specialized, so should the genera
be within a subfamily. While the concepts
of primitive and advanced are difficult to
apply with confidence to many of the fly-
catcher taxa, it should be possible to recog-
nize certain groups as generalized or spe-
cialized. If we accept the concept of the
manakins and flycatchers as evolving from
a common stem, then some of the small
green flycatchers of the restricted subfamily
Elaeniinae are probably closest to the an-
cestral stock. They are generally olive
green above, with the crown sometimes
gray or dusky; paler below, whitish, yellow
or olive; t^vo pale wing-bars and pale
edgings on remiges; bill short and con-
stricted, and light rictal bristles. Genera
that fall naturally into this group are
Xantliomyias, Phyllomyias, Tyronniscus,
Oreotriccus and Acrochordopus. They have
almost certainly di\erged some from the
common primitive stock l^etween manakins
and flycatchers, because that was presum-
ably a humid forest form as are many of the
manakins today, and the above genera are
found in drier, woodland habitats or in the

146 Bulletin Mu.scudi of Comparative Zoology, Vol. 148, No. 4

subtropical zone. They are, howexer, uii-
specialized, at least morphologically.

The characters used to define the above
genera are minor. PJiiilloniyias has a
broader, more triangular bill and heavier
rictal bristles than Xanthomiiia.s: Oreo-
tricctis is considered to have an exaspidean
tarsus while the others are pycnaspidean or
"quasi-exaspidean"; Tyranniscus has a
short, compressed bill; AcrocJiordopus has
the most pointed wing and a pycnaspidean
tarsus with the distal scntella roughened;
Oreotriccus has a rounded wing. Plumage
patterns, however, cut directly across the
generic lines. Acrochordopus Inirmeisteri,
PJu/Uomyia.s f(isci(ifiis\ and Xanthomyias
virescens are virtually identical in plumage;
they can only be distinguished by the
morphological characters listed above; the
same is true of the pairs Oreotriccus pluin-
heiceps and ^rip-anniscus cinereiceps, and
XatitJiODiyias .sclateri and Fhylloviyias
{Oreotriccus) ^riscocapilla. The last named
species, griseocupilla, was transferred from
Phyllomyias to Oreotriccus by Zimmer
( 1955: 23) because of its exaspidean tarsus.
However, Fhyllomyius fasciatus shows indi-
\'idual variation from exaspidean to taxas-
pidean (Zimmer called it pycnaspidean, but
with birds this size the tarsal types are dif-
ficult to distinguish), and the tarsal scu-
tellation is not a valid generic character.
With the exception of a group of species
now in Tyranniscus that I separate below,
I unite in one genus Xanthomyias, PJiyllo-
myias, Tyranniscus (nigrocapillus, uropy<s,i-
alis, and cinereiceps only ) , Oreotriccus and
Acrochordopus. PlnjUomyias and Tyran-
niscus are e(|ually available (both de-
scribed by C^abanis and Heine, 1859: 57) as
the oldest name; I select Plujllomyias since
I split the present genus Tyranniscus, and
the use of that name might cause confusion.

Zimmer (1955: 24) pointed out that the
species of Tyranniscus fall into two groups
on wing pattern. The species ni<i,rocapillus
(type of genus), uropygialis and cinereiceps
have the common wing pattern of two
wing-bars, pale edgings on the flight

leathers, and a contrasting black patch at
the base of the secondaries. The remaining
species, vilissimus, bolivianus, cinereocapil-
lus, iiracHipes and viridiflavus, have a pat-
tern unique among the tyrannids. The
median and greater coverts are edged along
the outer webs with yellow or white, rather
than having the pale coloring at the tips
producing a wing-bar; the secondaries and
four or five subexternal primaries are edged
with yellow, but the outermost and the
four or five inner primaries are black. This
produces a distinctive black stripe (wedge)
down the wing when it is spread. Corre-
lated with this type of wing is a longer bill
and a much less accentuated black specu-
lum on the secondaries. Zimmer believed
both types had (juasipycnaspidean tarsi,
but in my notes on the tarsi, made before
I was aware of the plumage types, I called
ni<irocapillus, uropyfi,iaIis and cinereiceps
"pycnaspidean /taxaspidean" and vilissimus,
bolivianus, cinereocapillus, ^racilipes and
viridiflavus "exaspidean, some pycnas-
pidean proximally." Ames (1971: 162)
noted a distinction in the syrinx. He found
)ii^rocapiUus like OrnitJiion, and cJirysops
(a race of viridiflavus) like nothing else.
His comment (p. 162) was, "The two spe-
cies of Tyranniscus examined differ so strik-
ingly in syringeal moiphology that I felt it
necessary to verify the identity of the speci-
mens through comparison with skins. One
cannot help wondering if a thorough analy-
sis of structural and behavioral characters
of these and other species of Tyranniscus
would not result in dividing the genus."
Warter (1965; 157) examined specimens
of Tyra)iniscus acer (a race of iiracilipes),
chrysops and vilissimus. They had the type
6 nasal septum, otherwise found only in the
Flnvicolinae and the aberrant Polystictus.
Xantfiomyias virescens, the only other
member of my Phyllomyias group that he
examined, had a t\'pe 1 nasal septum, also
foimd in se\'eral other genera of the en-
larged Elaeniinae.

Published behavioral data on Tyrannis-
cus are virtually non-existent. However,

Classification of Tyrant Flycatchers • Traylor 147

John Weske {in lift.) says niiirocapillu.s
forages like a kinglet, but hoUvianus
perches and sallies like a typical flycatcher.
Distributionally the tliree barred-wing
forms, niii,wc(ij)illus ct al., are all Andean
subtropical; the "edged" forms are divided,
vilissimus occurring in Central America,
and Caribbean Colombia and \^enezuela,
and gracilipes in Amazonia, wliile J)oIivi-
anus, cinereicapilliis and viiidiflavus are
found in the Andean subtropics, sympatric
with the ni<!,wcapillus group. In the Vilca-
bamba Range in Peru, Weske ( in lift. )
found the range of the edged-winged
bolivianus overlapped those of the two
barred-winged species nigrocapiUiis and
uropij'gialis, while tlie last two replaced
each other altitudinally without overlap.
Only two nests have been recorded —
Snethlage (1935: 53S) reported the nest of
acer, a race of <i,raciUpes\ as large, triangu-
lar, retort-shaped and hung from the tree
branches at middle height, and Skutch
(1960: 465) described that of vilissimiis as
a cozy ovoid structure with a side doorway,
found in hanging mosses or bromeliads.
Unfortunatelv, ocer and vilissimus have the
same wing type and there is no compara-
tive material of ni<j:,rocapiUus et al. FhijUo-
mijias g^riseocapiUa and biirrneisteri, how-
ever, both build cup-shaped nests.

Despite the superficial similarity, I be-
lieve the group of Tyranniscus species with
the unique wing pattern, syrinx and aber-
rant nasal septum, should be separated
generically from the typical group of spe-
cies. Surprisingly, there is no name avail-
able, so I propose:

Zimmerius gen. nov., type Tyrannulus
chrysops Sclater, 1858.

Diagnosis. Superficially most like Tyran-
niscus Cabanis and Heine, 1859, but differs
as follows: has a unique wing pattern in
which the median and greater coverts, the
secondaries and four or five subexternal
primaries are edged with yellow, while the
outermost and f(nu- or five internal pri-
maries are blackish, producing a black

wedge when the wing is spread; tail pro-
portionately longer, greater than 80 per
cent of wing length instead of less than 80
per cent, and bill proportionately longer,
greater than 18 per cent of wing length;
tarsus basically exaspidean with a few
scutes on the proximal half of the plantar
surface, compared to taxaspidean/pycnas-
pidean in Tyranniscus; syrinx unlike any
others in the Elaeniinae, lacking intrinsic
muscles; nasal septum of Winter's (1965:
34) type 6 with transverse plate, a type
found elsewhere only in the Fluvicolinae
and in the unrelated Polystictus of the
Elaeniinae. When Tyranniscus is merged
with Fhyllomyias (inch Xanthomyias and
Oreotriccus) the wing pattern, syrinx and
nasal septum of Zimmerius remain as diag-
nostic characters, but the different propor-
tions and tarsal envelopes are covered by
variation in PJtyllomyias.

The species included in Zimmerius, all
formerly in Tyranniscus, are boliviaiuis,
vilissimus, cinereicapiUus, gracilipes and
viridiflavus (including chrysops). The
genus is named for the late John Todd
Zimmer in belated recognition of his enor-
mous contribution to the systematics of
Neotropical birds, and his recognition of
the unique wing pattern in this new genus.

Although I have Zimmerius follow Phyl-
lomyias in the lineal list, I am not con-
\inced they are closely related. There is,
however, no other genus to which they
seem more nearly related, so leaving them
together will at least retain the benefit of
familiarity.

Ornithion (including Microiriccus) is a
genus of (|uite small flycatchers with dis-
proportionately short tails. In plumage
pattern, O. inerme is not unlike Phyllo-
myias (Tyranniscus) nigrocapillus, but
much smaller; the other two species lack any
wing-bars. Ornithion inerme is intermediate
in tail length between ""Microiriccus" hrtin-
neicapiUum and semifhivus and the shortest
tailed PhyUomyias in its size range. The
values of the tail,' wing ratios are: hrun-
neicapiUum and semiflavus, 54 per cent and

148 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

57 per cent; inernie, 70 per cent; and P.
grisciceps, 87 per cent respectively. The
tarsi are taxaspidean/pycnaspidean, but the
tarsi are so variable in this whole Elaeniine
group of flycatchers, that they are not good
indicators of relationship. Ames found the
syrinx of semiflavus to be near P. niiiro-
capiUus and probably related to Elaenia.
In cranial characters Ornithion seems
typical of many small flycatchers. Although
there are no really trenchant characters to
diagnose Ornithion, the included species
are a cohesive group, and I recognize it.

Tymnnidiis elatus is a small genus and
species, similar to the smaller Zimmeriiis in
size and proportions, but nearer Phyllo-
mijias nigrocapiUus in plumage pattern. It
has generally been included with these
genera in lineal hsts, but it differs from
both of the above in having a partially
concealed bright yellow crest. Bright crests
are common in the flycatchers, and have
midoubtedly arisen independently in a
number of different lineages. However,
they are not known to occur in only one
species of an otherwise crestless genus, and
I do not think that elatm- belongs in either
PhijUomijias or Zimnierius. On the other
hand, elatus is almost a miniature of Myio-
pagis gaimardii, and I believe that is where
its relationsliips lie. Both genera have
taxaspidean tarsi, and Ames (1971: 161)
included T ijrannulus and Mijiopagis in his
Elaenia group on syringeal characters.
Warter (1965: 34), however, found that T.
elatus and M. gainiardii (which he included
in Elaenia) differed in their types of nasal
septum, interorbital septum and cranium.
The relationship of elatus with M. viridi-
eata was closer, but both skulls were
damaged, so that the evidence was incon-
clusive. Tyrannulus should be placed next
to but not in Myiopagis.

Caniptostoma, Phaeomyias, Suhlegatus,
Suiriri, Myiopagis and Elaenia are a group
of closely related genera. They are more
successful than the genera previously dis-
cussed in the sense that all except Suiriri
are found throughout the tropical lowlands

of South America, and three reach Mexico
or the scnith western United States. With
the exception of Suhlegatus, all are in-
cluded in Ames' Elaenia group on syringeal
characters.

Phaeomyias is basically a monotypic
genus for the type, murina, but Hellmayr
and Zimmer added or subtracted other spe-
cies. Hellmayr (1927: 453) included
tenuirostris, a species that Zimmer (1955:
2) properly removed to Ineziu. Prior to
that Zimmer (1941b: 10) added EUenia
leiicospodia because of its partially taxas-
pidean tarsus. The latter character, how-
ever, is of little value by itself; in some
species, such as Phyllomyias fasciatus, the
tarsi can vary from taxaspidean to exas-
pidean with every gradation in between.
Leucospodia has a partially concealed
white crest, similar to those of Elaenia or
Myiopagis, and should not be placed in a
crestless genus without more substantial
evidence. Actually, in every available
character of size, proportions, pattern and
nest-type, leucospodia is a typical Myio-
pagis; the only difference is that all Myio-
pagis have at least some yellow or olive in
their plumage (as does Phaeonujias murina),
while leucospodia is plain dull brown,
whitish below. This latter coloration is not
unusual in species endemic to the arid coast
of Peru, and I place leucospodia in Myio-
pagis.

Pluieonujias murina is about as "typical"
a dull brownish flycatcher with yellowish
belly as one could visualize; its only dis-
tinctive external character is a fairly broad,
whitish superciliary. Camptostonm is a
smaller version of the same, but with a
compressed, arched bill. Both are mem-
bers of Ames' Elaenia group, along with
Suiriri, Myiopagis and Elaenia, but Phaeo-
myias and Camptostoma have type 2 nasal
septa, without the transverse plate, while
Suiriri and Elaenia have type 5 with plate.
The situation in Myiopagis is uncertain,
with gainiardii having type 5, viridicata
pr()l)ably type 1, and the others not ex-
amined.

Classification of Tyrant Flycatchers • Traylor 145

Camptostoma and Phaeomyias agree in
general form and color, and in all syringeal
and most cranial characters; however, they
differ strongly in voice and behavior ac-
cording to both Eisenmann and Fitzpatrick
(in litt.). One of the most marked differ-
ences between them is in the form of their
nests. Camptostoma builds a globular nest
with a side entrance, placed in branches
of low trees, while PJuieomyias builds a
neat cup, typical of the Elaenia group of
genera. Ihering considered the nest of
Camptostoma transitional between the cup-
shaped nest of his Elaeniinae and Serpo-
phaginae, and the pendant nest of his Eus-
carthminae. However, Camptostoma itself
shows no relation to the Euscarthmines,
l)eing a typical member of the Elaenia
group in color, form and syringeal char-
acters. Despite its close resemblance to
Phaeomijias, I keep it separate because of
the differences in behavior and nest form.

The genus Suhlegatus is composed of
three parapatric species ( to be discussed in
a separate paper) found throughout prac-
tically all of lowland South America, al-
though probably not resident in the rain
forest. It differs from most Elaenia only in
the lack of a white crest, and its somewhat
swollen bill with wholly black mandible; it
can be told from E. cristata only by the bill
characters. It builds a shallow cup-shaped
nest like Elaenia, and Warter (1965: 34)
records them as having similar, but not
identical, crania. In fact, there is little be-
sides intuition that leads me to keep Suh-
legatus out of Elaenia, and the fact that the
three species of Suhle<iatus form a different
speciation pattern from that of any Elaenia.

Suiriri is composed of two hybridizing
taxa, which Zimmer (1955: 18) thought
should be treated as a single species, but
which Short (1975: 283), presumably
working with the same material, maintains
as two species. They differ from Elaenia
in lacking a white crest, and in having a
longer bill with solid black mandible. The
eastern species affinis differs from any
Elaenia in having the rump and proximal

quarter of the rectrices pale yellow in con-
trast to the olive back and blackish tail;
however, it differs from the congeneric
(possibly conspecific) suiriri in the same
way. The most distinctive character shown
by Suiriri is the white spotting of tlie
Juvenal plumage. All the grayish-brown
dorsal feathers of the juvenal plumage have
a wedge-shaped white tip, as do the wing
coverts and three inner secondaries. When
viewed from above, the juvenal bird shows
a startling resemblance to juvenal Old
World flycatchers of the genus Muscicapa;
the pattern is unique in the Tyrannidae,
except that juvenal Siihlegatus has white
spotting on the crown, according to Fitz-
patrick (in litt.).

Although Myiopagis has always been
associated with Elaenia, and many authors
have followed Hellmayr (1927: 401) in
uniting the two, Zimmer (1941a: 20) con-
sidered them distinct genera. He kept them
separate because of the pycnaspidean or
taxaspidean tarsus of Myiopagis, and the
usually yellow ( white in some subspecies of
gaimardii) rather than the usually white
crown in Elaenia. Ames found the syrinxes
alike, but Warter (1965: 37) recorded M.
viriclicata ( the type of the genus ) as having
a type 2 nasal septum, while gaimardii had
a type 5 like Elaenia. As pointed out by
Eisenmann (in litt.), Elaenia species prefer
open habitats where they are relatively con-
spicuous, and Myiopagis species prefer
denser foliage where they are relatively in-
conspicuous. I recognize Myiopagis, but
with the knowledge that further anatomical
research may suggest either merging it
with Elaenia, or transferring species be-
tween them.

Elaenia is the largest and most "success-
ful" of the Elaeniine genera, with 19 spe-
cies occurring from southern Mexico to
Tierra del Fuego, and from both humid and
arid lowlands to above 3000 meters in the
Andes. Although there is no single charac-
ter by which to diagnose the genus, it is a
natural assemblage whose limits are gener-
ally accepted. In size, the species range

150 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

from medium to large for this group of
genera, wings 70 mm to 91 mm, with rela-
tively short bills, 16-18 per cent of wing
except for crisiata in which it is 20 per cent
and short to medium tarsi, 20-26 per cent
of wing. The bill is short, moderately
broad at the base and deep throughout,
rictus lightly bristled, and wing usually
fairly pointed, 10th primary ecjual to 4th to
6th. Plumage is generally dull olive above,
except for males of strepera, which are dark
slate gray, and the undeiparts are pale yel-
lowish to ohve to whitish; the majority of
species have a concealed white crest. As far
as recorded, all species build a neat cup-
shaped nest, frequently covered with
lichens.

The genera thus far discussed make up
Warter's tribe Elaeniini, with the exception
of Leptopogon, Mionectes and Pipro-
morphch which will be inserted in the lineal
list further on. The adjoining tribe of War-
ter, the Serpophagini, I consider to merge
into the Elaeniini without the slightest
break. Serpophciiia and Myiopagis have
been confused in the past, M. caniceps hav-
ing been described in l^oth genera, and as
recently as 1957 Dorst described a popula-
tion of Myiopagis gaimardii as Serpophaga
herliozi (see Mayr, 1971: 313). Mecocer-
cithis is also an obvious bridge. Mecocer-
ciiJus species are remarkably close in plum-
age to FhyUomyias species. Intergeneric
pairs showing especially close resemblance
are M. hellmayri and P. iiwpygialis, M.
minor and P. phimbeiceps, and M. poecilo-
cercus and P. sclateri. The only plumage
character that consistently distinguishes
Mecocercuhis is the distinct white super-
ciliary. On the other hand, Ames (1971:
73, 74) found the syrinx of Mecocerculus
like that of Serpophaga and Anairetes.
Smith (1971: 285), on the basis of his be-
havioral studies of Hellmayr's Seqoophag-
inae, considered Mecocerculus closely re-
lated to Serpophaga (inch Inezia), Anairetes
(inch Uromyias) and Stignwtura. Finally
Warter (1965: 34) recorded Mecocercuhis
as having a type 5 nasal septum, which it

shared only with Elaenia and its closest
relatives, and with Serpophaga, Inezia, and
Anairetes. There seems little question that
the foiTner subfamilies or tribes were arti-
facts.

The genus Inezia is similar to Serpo-
phaga in size, proportions and general
plumage pattern. Hellmayrs Inezia con-
sisted only of the species siibflava, but Zim-
mer (1955: 1) added to it Serpophaga
inornata and Phaeomyias tenuirostris.
Smith (1971: 266) included Inezia in Ser-
pophaga because of similarities in appear-
ance and behavior. However, Parkes (1973:
249 ) has pointed out that the Inezia species
differ from Serpophaga in lacking a white
crest, and in having taxaspidean instead of
exaspidean tarsi, pale instead of black
mandibles, and a distinctive juvenal plum-
age rather than one like the adult plumage.
I consider this combination of characters
sufficient to define the two genera. There
is also a close resemblance in plumage be-
tween Inezia snbflava and the two species
of Stigmatura. The latter are characterized
by greater size, proportionately long tails
and a nasal septum without a basal plate.
The relative tail lengths compared to wing
lengths of Stigmatura budytoides and mi-
pensis are 121 per cent and 113 per cent
respectively, compared to 98 per cent in
.sul)flava. The plumage pattern of the three
species is basically the same, uppeiparts
olive brown, with a prominent white or
pale yellow superciliary stripe running to
well behind the eye; underparts pale yel-
lowish; two distinct white wing-bars, and
flight feathers edged white. The tails of
both are rounded, but those of budytoides
and napensis are elaborately patterned
with white, while that of snbflava is only
narrowly tipped. Although the resemblance
is close, I recognize Inezia and Stigmatura
as distinct genera, primarily because of the
difference in nasal septa, the former having
a type 5 and the latter a type 2.

Anairetes, Uronujias and Yanacea are a
closely related group of genera. They are
composed of long-tailed, long-legged spe-

Classification of Tyrant Flycatchers • Traylor ISl

cies, found in tlie upper subtropical and
temperate zones of the Andes and south
through Chile and Patagonia to Tierra del
Fuego. Yanacea alpinus and Uromyias
a<i,raphia are practically unstreaked, recall-
ing Stigniatura, but Uromyias agilis is
heavily streaked on throat and breast, simi-
lar to Anairetes species. Uromyias was
separated from Anairetes because of its
long tail with pointed rectrices, and Yana-
cea was recognized because of its relatively
shorter bill and tarsus. These characters do
not seem of great importance in an other-
wise closely related group, and I merge
Yanacea and Uromyias in Anairetes. Zim-
mer (Ms.) had already merged Yanacea
and Anairetes, as he suggested in an earlier
discussion (1940b: 10); Smith (1971: 275)
merged Uromyias and Anairetes.

Tachtiris, the brilliantly colored Siete
Colores, has regularly been associated with
Serpophag,a. Smith (1971: 2S4) says it
probably belongs with the Serpophagines,
but it may have closer relatives in the Eus-
carthmines. He did not specify which of
the latter, but possibly it is near Pseudo-
colopteryx, which also has marsh-living
forms. However, the color and patterning
of Tachuris are unique, particularly the
blue auriculars, and there is no way to re-
late it closely to any of the other smaller
flycatchers. The combination of orbital
septum, palatines and cranial type are
found throughout the Elaeniinae, and the
nasal septum was undetenuined (Warter
1965: 34); Ames found the syrinx unlike
any other. I shall leax'e Tachtiris in the
vicinity of Serpopluiga, but it could equally
well be sedis inceiiae.

Colorhamphus parvirostris was placed in
the genus Ochthoeca by Berlepsch (1907:
470), but was kept in its monotypic genus
near Serpophaga by Ridgway (1907: 396)
and Hellmayr (1927: 400). I agree with
Berlepsch, and discuss parvirostris in more
detail under Ochthoeca.

CuUcivora, Pohjstictus, Pseudocolopteryx
and Euscarthmus have been kept at the end
of the Euscarthminae since Berlepsch

( 1907 ) . The subfamily Euscarthminae was
created by Ihering ( 1904 ) for a group of
genera making pendant, pyriform nests,
and its type genus Euscarthmus had at that
time for its type species Euscarthmus nidi-
penduhis W'ied, a species now in the genus
Idioptilon. Culicivora and allies show no
particular resemblance to the typical Todi-
rostrum/ Idioptilon group, and are in some
ways nearer Serpophaga. All but Euscarth-
tnus have one or more species with whitish
or streaked crests as in Serpophaga, they all
have weak rictal bristles, and all make cup-
shaped nests. In bill length they are inter-
mediate between the long-billed Todiros-
trum and the shorter-billed Serpophagas.
They seem to form a natural group, with
their center of distribution in the dryer
country of southeastern Brazil and northern
Argentina and, except for Culicivora, with
isolated populations in dry country north
of the Amazonian forest. Pseudocolopteryx
has penetrated the more temperate zones
in Patagonia and the southern Andes, and
three of the four species are sympatric in
the Chaco. The four genera seem worth
recognizing. Culicivora has a most peculiar
rounded tail, with only 10 rectrices, and the
barbs are stiff and decomposed, recalling
Synallaxis in the Furnariidae. Polystictus
has an aberrant cranium, with nasal sep-
tum, interorbital septum and palatines
characteristic of Fluvicola rather than the
Elaeniinae. Pseudocolopteryx is a natural
assemblage of predominantly yellow spe-
cies with marked sympatry among them,
suggesting a long phylogenetic history;
three of the four species have aberrant pri-
maries. The two species of Euscarthmus are
rich browns and buffs with rufous crests;
they have frequently been allied to Pseudo-
colopteryx, but I hesitate to unite them be-
cause the latter is so uniform without Eus-
carthmus.

Leptopogon, Mionectes and Pipromorpha
were placed at the end of the restricted
Elaeniinae by Berlepsch (1907: 492). Pip-
romorpha had been merged in Mionectes
for many years until Ridgway (1907: 354)

152 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

WING LENGTH

100-

95-

5 90-

85-

80-

venezuelensis

angustirostris

difficilis
X

superciliarls '-'

Capsiempis

qualaquizae

paulistus

Leptotrjccus

flavovirens
X

virescens

oustaleti
X

ophthalmicus

I
poecilotis

nigrifrons

chapmani

Figure 1. Proportions of tail/wing plotted as functions of wing length for species of the genera Phyllos-
caries, Pogonotriccus, Leptotriccus and Capsiempis. Although these genera were originally separated because
of differing proportions, the points fall into a continuum. Closed circles, "•", are Pogonotriccus, crosses,
"X", are Ptiylloscartes, and open circles, "O", are the types of the monotypic genera Leptotriccus and Cap-
siempis.

resurrected it l^ecaiise of the different
.shapes of the 9th primaries in adult males.
This is a trivial character in a family where,
in a single genus such as P.seudocolopterijx,
three species have different sets of pri-
maries aberrant, and the fourth has them all
normal. In all other morphological and
anatomical characters — size and propor-
tions, bill shape and lack of rictal bristles,
syrinx and cranial characters — Pipro-
morpha and Mionectes are virtually identi-
cal, and I again merge them. Leptopog,on
is close to Mionectes in size and propor-
tions except for its longer tail, and both
genera make pendent, globular nests, usu-
ally hung under logs or cut banks in the
vicinity of water; this is unlike the nests of
any Elaeniine flycatcher. According to
Monroe ( 1975 ) , they also share the be-
havioral trait, unusual among the Tyran-
nids, of single-wing flicking. Besides the

peculiar nest, Leptopogon differs from the
Elaeniine group in having fairly heavy
rictal bristles, but this character is not
shared with Mionectes. Leptopogon differs
from Mionectes in having a much more
rounded wing, longer tail, heavier rictal
bristles, and normally shaped primaries; in
all Mionectes species but nifiventris, males
have either the 9th or 10th primary notched,
shortened or attenuated. According to pub-
lished accounts, there seems to be little
difference in behavior.

The section of the subfamily where Lep-
topogon and Mionectes seem most at home
is with Plnjlloscartes, Pogonotriccus and
allies. In plumage and pattern, Leptopogon
is particularly close to PJiylloscartes and
Pogonotriccus, in fact four of the species in
the latter genera were originally described
in Leptopogon. Superficially, there is little
difference between Leptopogon and PJujl-

Classification of Tyrant Flycatchers • Tmylor

25-

24-

23-

uj 22-

21 ■

20-

19-

WINU LENGTH

52 54 56

flavivenlris
venezuelensis

pauhstus

qualaquizae

superciliaris

ventrahs
X

angustirostris
X

Capsiempis i difficilis

f lavovirens

Leptotriccus

eximius

• orbitalis

oustaletl

virescens

ophthalmicus

nigrifrons

chapmani

poecilotis

Figure 2. Proportions of culmen, wing plotted as functions of wing length. (See Fig. 1. legend.

loscartes, but the former has a shorter
tarsus, a type 3 nasal septum found other-
wise only in the unrelated genera Plotijiin-
chiis and Omjchorhiinchus, and the peculiar
nest type and single-wing flicking men-
tioned above. The importance of the nest
type cannot be evaluated, since the nest
type of only one of the 17 species in Po-
gonotriccus and PhyUoscartes is known;
PhyJJoscartes venfralis builds a partially
domed, somewhat globular structure in the
fork of a bough, according to Ihering (1904:
314).

PhijUoscartes, Pogonotriccus and the two
monotypic genera Leptotriccus and Cop-
siempis are a closely related group of
genera placed by Berlepsch and Hellmayr
in the Euscarthminae. They are all similar
in appearance — small greenish and yellow-
ish flycatchers, mostly with two well
marked wing-bars, and several with gray

crowns and patterned faces. All four were
described as monotypic genera by Cabanis
and Heine ( 1859 : 52-56 ) for the type spe-
cies PJujUoscartes ventraUs, Pogonotriccus
eximius, Leptotriccus syJvioIus and Cap-
siempis flaveola. The first three were de-
fined primarily on mensural characters;
PhyUoscartes with a long bill and tarsus,
Pogonotriccus with a long wing and short
bill, and Leptotriccus by a "Serpophaga-
like" bill and long tail. Capsiempis was
considered more like Tolmonujias flavi-
venter, with a broad bill. During the next
hundred years 15 species were added to
this complex, seven by original description
and eight by transfer from other genera,
primarily Leptopogon and MecocercuJus.
Within the group, two species, originally
described in Leptotriccus and Capsiempis
respectively, were moved to Pogonotricctts.
\Mien the proportions of tail, culmen and

154 BuUetin Miiscinu of Comparative Zoology, Vol 148, No. 4

WING LENGTH

36-

34-

32-

30-

28-

26-

24-

llaviventns

paulistus

supercilians
X

qualaquizae

venezuetensis

ventralis

Capsiempis

Leptotnccus

flavovirens

angu5tirostris

ditficilis

orbitalis

vtrescens
X

oustaleti
X

poecilotis

nigrifrons

chapmani
X

ophthalmicus

Figure 3. Proportions of tarsus wing plotted as functions of wing length. (See Fig. 1. legend.)

tarsus to wing length are plotted as func-
tions of wing length (Figs. 1-3), it is clear
that on mensural characters the present
genera overlap widely. There is a general
trend within the group for species with
longer wings to have proportionately
shorter bills and tarsi, but this is true of all
the currently recognized taxa. Even though
some species are obviously separated from
their nearest relatives by the present allo-
cation to genera — the representative spe-
cies chapmani and poecilotis being in
Phylloscartes- and Pogonotriccus respec-
tively— no shifting of species between
genera would segregate out two or more
taxa. Warter (1965: 33) found the cranial
characters similar in all four genera. Ames
(1971: 67-72) considered the syrinxes
different from each other and from all other
genera, but as noted before negative syrin-
geal evidence is inconclusive. I unite the

four genera into a single genus. All the
names were introduced at the same time in
the same pubHcation, and are equally avail-
able under the Rules [Art. 24(a)]. I select
Phylloscartes as the name for the combined
genus.

The geographical distributions of the spe-
cies are most peculiar, but shed little light
on relationships. Five species of Pogono-
triccus are found in the subtropics of the
Andes from Venezuela to Bolivia, but the
sixth, the type eximius, is confined to the
southeastern forest region of Brazil and
adjoining Paraguay and Misiones, Argen-
tina. On the other hand, four Phylloscartes
species and Leptotricciis sylviolus are en-
demic to the southeastern forest region, and
only the type, ventralis, has an isolated
population in the Andean subtropics. The
remaining species of Phylloscartes are
found in the Guianan forests, the tepuis of

Classification of Tyrant Flycatchers • Traylor

Venezuela, and in eastern Central America.
Despite the fact that these are all forest
forms, no species of this complex occurs in
the Amazonian forests.

The one species that is out of place on
ecological grounds is Copsiempis flaveolus,
which has adapted to shrubs and bushes
in clearings, and is found in the drier parts
of tropical South America, north and west
to Nicaragua. Eisenmann (in litt.) says
that where Capsiempis overlaps the range
of Phylloscartes flavovirens in Panama, the
two are unlike in habits and do not appear
closely related. However, as Zimmer
(1940b: 2) pointed out, flavovirem and the
closely related virescens are unusual among
the Phijlloscai'tes species in having more
pointed \\dngs, with the 10th primary equal
to or greater than the 4th, rather than ef{ual
to or less than the 1st. Pog^onotriccus spe-
cies and Leptotricctis also have rounded
wings with a short 10th primary. Prac-
tically nothing has been published about
the habits of any PhijUoscaries or Pofi^ono-
triccus, so it is not possible to say if flavo-
virem is typical of the rest of the genus.
The only nests that have been described are
flaveolus nests, which are cup-shaped, and
ventralis nests which are partially covered.
Although Capsiempis may later prove
worthy of reccjgnition as a distinct genus,
I keep it in the enlarged PlujUoscartes on
morphological grounds.

Although the species of Phylloscartes do
not have the long, spatulate-type bill char-
acteristic of Todirostriim and Idioptilon,
the bill is proportionately longer, and the
rictal bristles more developed, than in the
genera discussed so far. Their plumage
pattern is typical of the "small green fly-
catchers" such as Pliyllomyias and Meco-
cerculus, and they with Leptopo<^on repre-
sent the primitive stock from which the
Todirostriim group evolved. Figiue 4 is a
schematic diagram showing the probable
origin of the three best defined lineages
within the Elaeniinae — the Elaenia group,
the Tit-tyrants (Anairetes) and the Tody-
tyrants. The position of Pseudocolopteryx

PSEUDOCOLOPTERYX el al

TIT-TYRANTS '

/ PSEUDOTRICCUS

/ /'

TODY-TYRANTS

ELAENIA GROUP

SERPOPHAGA

PLATYRINCHUS el al.

PHYLLOSCARTES

LEPTOPOGON
/
VMECOCERCULUS/'

PHYLLOMYIAS |

I /
1/

Figure 4. Schematic diagram showing probable re-
lationships among the Elaeniinae. The three clearest
lineages, Elaenia group, Tit-tyrants and Tody-tyrants,
arose from Phyllomyias. Mecocerculus. and Lepto-
pogon and Phylloscartes, respectively, genera that
share a basic, "little green flycatcher" plumage pat-
tern. The positions of genera not part of these
lineages can only be suggested.

and allies, and of Pseudotriccus is probably
somewhere between the Tit-tyrants and the
Tody-tyrants, although not part of the
lineage of either; the position of the Flat-
bills, Platyrinchus and allies, is even less
clear. PhyUomyias, Mecocerculus, and
Leptopogon and PJiylloscaiies, the presum-
ably primitive genera, resemble each other
closely, although they lead into three
distinct groups.

Pseudotriccus, including Caenotriccus
ruficeps (Zimmer, 1940a: 22), is a peculiar
genus of three species, confined to the sub-
tropics of the Andes and extreme eastern
Panama, that seems to have no close rela-
tives among the genra of Hellmayr's Eus-
carthminae. It has generally been asso-
ciated with Hemitriccus at the end of the
Todirostrum group, but there is little real
resemblance. The colors of Pseudotriccus
are dull olive, olive brown or rufous, and
the birds are without any embellishments
such as streaking, superciliary stripes, eye
rings, wing-bars or contrasting edgings to
the flight feathers. All of the Todirostrum
group and Phylloscartes have yellow under
wing coverts that come around the edge of

156 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

the wing, giving a bright spot at the bend
when the wing is folded. In Pseudotriccu.'i
the under wing coverts are Hke the body
phmiage, \\'\\h no contrast. The one genus
to wliich P.scudotriccus may be alHed is
Corytliopis, which has only recently been
admitted to the Tyrannidae and which is
without obvious relati\'es. The two genera
share a proportionately long tarsus, rela-
tively as long as those of the much smaller
Todirostrum/Idioptilon species; the scutes
of the tarsi are almost obsolete, giving a
smooth booted effect; the wings are plain
without any pale bars or edgings; and the
rectrices are broad and soft. Fitzpatrick
(in conversation) says they share similar
feeding habits, walking along the ground
and leaping up to pick insects from the
under sides of leaves. I place Conjthopis
and Pseudotriccus together, and leaxe them
between PlujUoscarte.s and the Todiro.stmm
group of genera, not because I am con-
vinced that is where they belong, but be-
cause I know of no better place.

The genera I include in the term
^^Todiro.stmm group," as used above, are
the remaining genera of Hellmayr's Eus-
carthminae: Hemitriccu.s, Peri.s.sotricciis,
Myiornis, Atalotriccu.s; Colopteryx, Lopho-
triccus, Idioptilon, Taeniotriccus, Poecilo-
triccu.s, Snethliiiiaea, Microcochleariu.s,
Euscarth mornis, Onco.stotna, Ceratotriccns
and Todiro.stmm. They are a closely related
group, characterized by long slender tarsi,
greater than 29 per cent of wing length,
and long wide bills, greater than 25 per
cent of wing length, culminating in the
spatulate bill of Todirostmm. A general-
ized plumage pattern is found in at least
some species of most genera: olive green or
brown above; whitish below on throat and
breast with darker streaking, and yellowish
or olive on abdomen; remiges edged with
olive or yellow. All species have yellow
imder wing coverts, which usually extend
enough around the bend of the wing to
make a yellow spot when the wing is
folded. All recorded nests are of the pen-
dent, purse-shaped or pyriform type.

Of the genera that Ames (1971: 67)
examined all belonged in his Colopteryx
group except Todiro.stmm. The cranial
characters were alike in those genera that
VVarter (1965: 33) examined, but similar
crania were found in PJiyllo.scartes and
Pseudocolopteryx, and in Rhynchocyclus,
Tolmomi/ia.s and Cnipodectes of Hellmayr's
Platyrinchinae. Zinmier (1940a: 13, 22)
merged EuscartJimorni.s into Idioptilon, and
Perissotriccus into Myiornis, and these
changes were accepted by Meyer de
Schauensee ( 1966 ) .

Myiornis (including Perissotriccus) is a
genus characterized by minute size; wing
length in males averages less than 40 mm in
all three species, and they are the smallest
of the Tyrannids. Tail length is very short
in ecaudattis, the type of Perissotriccus, but
it is normal in auricularis and intermediate
in alhiventris. Bill and tarsal proportions
fall into the normal range of the tody-ty-
rants.

Lophotriccus, Colopteryx and Atalo-
triccus have been placed together in the
past, but the evidence for including Atalo-
triccus is equivocal. Lophotriccus is dis-
tinguished from the remainder of the tody-
tyrants by its distinctive crest; the crown
feathers are elongated and edged in con-
trasting gray or rusty. Colopteryx galeatus
is the same as Lophotriccus in plumage, but
has the three outer primaries much re-
duced. Atalotriccus pilaris has a normal
crown, but has the four outer primaries
even more reduced, narrowed and pointed.
The plumage patterns and crest of Lopho-
triccus and Colopteryx are so much alike
that the relationship must be close. I merge
them here because I do not consider
peculiar primaries a "generic" character in
the flycatchers. Atalotriccus has been kept
with Colopteryx because both have short-
ened outer primaries. If Atalotriccus has
indeed been derived from C. galeatus, then
the ornamental crest must have been lost
secondarily. But it is just as easy to con-
ceive Atalotriccus as an Idioptilon that has
independently acquired reduced outer pri-

Classification of Tyrant Flycatchers • Trayhr 157

maries and converged on Coloptery.x in this
character. The relatively long Atalotriccus
tarsns is more like that of hlioptilon than
that of Lophotriccus. Eisenniann {in litt.)
believes Atalotriccus behaves more like
Todirostrum sylvia than like Lophotriccus.
Considering the donbts about the origin of
Atalotriccus, I accept it as a distinct genus.
Since this tody-tyrant group is uniform in
cranial and syringeal characters, I doubt
if further anatomical study will help clarify
the relationships.

Fitzpatrick ( 1976 ) has just published a
valuable paper on Todirostrum and related
genera. He construed somewhat more
strictly the concept of "related genera," and
his study omits the following genera listed
above: Hemitriccus, Myiornis, Atalotriccus,
Colopteryx and Lophotriccus. Fitzpatrick
attempted to trace the lineage and generic
relationships of his tody-tyrants, and he
made the following taxonomic suggestions:

1. The genus Ceratotriccus should be
merged with Idioptilon.

2. The genera Taeniotriccus and Poecilo-
triccus should be moved to positions im-
mediately preceding Todirostrum, reflect-
ing their affinities with Todirostrum
capitale.

3. Todirostrum and Idioptilon should
continue to stand as separate genera.

4. The genera Snetldai^aea and Micro-
cochJearius are best merged with IdioptiJon^
while Oncostoma should continue to be
generically recognized.

Fitzpatrick's lineage is shown diagram-
matically in Figure 5, reprinted from his
Figure 4 (p. 443). The genera that he does
not consider, Myiornis, Atalotriccus, Colop-
teryx and Lophotriccus, would branch off
further down the stem, on the way to the
more highly specialized Todirostrum. \\^ith
the exception that I consider Hemitriccus
to belong to the "green" section of Idiopti-
lon, I believe Fitzpatrick's tree accurately
depicts the evolution and relationships of
the tody-tyrants.

If his diagram is correct, then his taxo-
nomic treatment of the copitale group, in-

'-- -">\ 1

1 Oncostoma

, Taeniotriccus / ^s,^
Poecilotriccus

THE

TODY -TYRANTS

Figure 5. Fitzpatrick's (1976, Fig. 4) tree showing the
lineage of the tody-tyrants. The stem to the lower left,
leading to Todirostrum capitale, is here considered
a single genus, Poecilotriccus.

eluding Todirostrum capitale, Taeniotriccus
andrei and Poecilotriccus ruficeps, must be
questioned. These three species are united
by a unique plumage pattern with chestnut
crown, and by the striking sexual dimorph-
ism in capitale and andrei. As Fitzpatrick
says (p. 441), "Affinities among the three
species are closer than those between the
lineage as a whole and Todirostrum or
Idioptdon." If this is so, the resemblance
between the bill of capitale and those of
Todirostrum species must be due to con-
vergence, and placing capitale in Todi-
rostrum makes that genus polyphyletic.
Fitzpatrick recognized that the three spe-
cies might possibly belong in a genus of
their own, but hesitated to make such a
major taxonomic shift without further
anatomical and behavioral study. How-
ever, I believe that the data require such a
move now, and I place capit(de, andrei and
ruficeps in a separate genus for which
Foecilotriccus is the oldest name.

Hemitriccus has been kept separate from
Idioptilon because of the former's allegedly
longer tail, and because of the slightly
shortened 7th primary in males. While tail
length in H. diops is proportionately longer
than that of any Idioptilon species, the

158 Bitlletin Museum of Comparative Zoology, Vol. 148, No. 4

same is not true of H. fkimmidatus whose
tail length is well within the normal Icliopti-
lo)i range. In males the 7th primary is
slightly shorter than the 6th and 8th, so the
edge of the opened wing shows a slight
notch rather than a smooth contour; the
female wing is normal. Considering the
variation in primaries within such genera
as Pseudocolopteryx, this characteristic
hardly rates generic separation. Plumage
characters are like those of Idiuptilun, and
female H. flammulatus can hardly be told
from the sympatric I. zosterops griseipec-
tus. Merging Hemitriccus with Idioptilon
will lead to yet another unfortunate change
in the name of this beleaguered genus.
Known as EuscartJiiniis to Sclater (1S88)
and Berlepsch (1907), it was Euscarthmor-
nis to Hellmayr (1927), Idioptilon to Meyer
de Schauensee (1966), and Cerototriccus to
Fitzpatrick (1976); it must now be called
Hemitriccus, the oldest available name.

To summarize the Todirostrum group,
the sequence of the genera and their synon-
ymies will be:

Myiornis (syn: Perissotriccus)

LopJwtriccus (syn: CoJopteryx)

Atalotriccus

Poecilotriccus (syn: Taeniotriccus)

Onco.stoma

Hemitriccus (syn: Idioptilon, Euscarth-

inornis, Snethlagaea, MicrococJdearius,

Ceratotriccus )
Todirostrum.

Hellmayr's Platyrinchinae — Platyrinchus,
Rhynchocyclus, ToJmomyias, Rompliotri-
gon and Cnipodectes — have occupied a
position between the "large" flycatchers
and the Todirostrum group since Berlepsch
( 1907), although Platyrinchus was put with
the Todirostrines by Berlepsch, and with
Rhynchocyclus et ah by Hellmayr. Warter
considered that on cranial characters the
Platyrinchinae graded into the Euscarth-
minae via Todirostrum and Oncostoma, but
in general form and appearance, they do
not seem closely related to any other genus
or group of genera. They have been kept

together in part because of their broad, flat
bills, moderate to heavy rictal bristles and
fairly short, slender tarsi, but in the case of
Platyrinchus, these are probably convergent
characters. The bills of Rhynchocyclus and
Tolmomyias are the extremes of broad and
flat but swollen-appearing bills with
strongly convex lateral edges; Cnipodectes
is somewhat narrower at the base, and
Ramphotrigon even more so, with the
lateral edges straight. Platyrinchus has the
same broad, convex bill, but it is much
flattened.

Rhynchocyclus and Tolmomyias re-
semble each other closely externally, the
latter seeming a smaller version of the for-
mer. However, Ames ( 1971 : 161 ) found
they "differ so strongly in their syringeal
structure that one may seriously question
the reliability of bill shape as a common
character." Also, Skutch (1960: 515) has
pointed out well-marked differences in
mating behavior, egg coloration and the
extent of down on the nestlings. Cnipo-
dectes is close to these two, but its brown
coloration, uniquely twisted primaries in
the male, and extreme sexual dimorphism
merit recognition. In plumage pattern,
Ramphotrigon species, particularly jusci-
cauda, look very much like Rynchocyclus,
but the bill is slender and straight edged,
the mandible is blackish rather than pale
horn (in dried skins), and they have two
well-marked buffy or chestnut wing-bars,
which are not found in Rhynchocyclus.
Ames (1971: 161) supported Zimmer's
(1939c: 16) transfer of megacephala from
Tolmomyias to Ramphotrigon. He found
the syrinx of megacephala strikingly differ-
ent from that of Tolmomyias sulphur escens.
The Ramphotrigon nest has not been de-
scribed; Rhynchocyclus and Tolmomyias
make pendent retort-shaped nests (Smithe,
1966: 322), and Cnipodectes makes a pen-
dent nest whose completed shape is not
known (Wetmore, 1972: 508).

As noted before, Platyrinchus does not
fit easily into this assemblage. The species
all have short tails, less than 50 per cent of

Classification of Tyrant Flycatchers • Traylor 155

ueij

wing length, a pale patch on the throat,
pale instead of blackish feet and tarsi and
brightly colored crests; they all build cup-
shaped rather than pendent nests. Pkitij-
rinchus has a type 3 nasal septum, found
elsewhere only in OmjchorJujnchiis and
Leptopogon, while Cnipodectes, Tolmo-
mijias and RhyncJwcychis have type 1.
However, in the last three, the nasal septum
evidently approaches the type 3, so the
difference may not be so great as it appears
at first glance. The syrinx of Platijrinclms
most resembles that of Ames' Colopterijx,
and is not at all like that of Tolmomyias or
Wiiinchocyclus. Although the relations of
Platyrinchus are almost certainly with the
Elaeniinae, it does not seem closely related
to any of the other genera, and I leave it at
the end of the subfamily.

FLUVICOLINAE

The subfamily Fluvicolinae as now con-
stituted, with the addition of the major part
of the old Myiarcliinae, has a number of
genera that in size and plumage pattern
resemble the more generalized Elaeniinae.
Such genera as Myiobms and Myiophohus
are close in size, proportions and plumage
pattern to the Elaeniine PhyJIomyia.s and
Phylloscartes. Externally they are best
distinguished by the broad, triangular
flattened bill and heavy rictal bristles;
internally they all have Waiter's type 6
nasal septum, which occurs elsewhere only
in Polystictus and Zimmerius, where it has
almost certainly been derived indepen-
dently. Within the FluvicoHnae there
appear to be two lineages. The first starts
with Myiohius, Myiophohus and allies, and
leads to Empidonax, Contopus and Sayor-
nis, which have speciated extensively in
North and Central America. The second
starts with the generalized Ochthoeca and
runs through the highly successful Xohnis
and Miiscisaxicola to a number of truly
aberrant genera such as Miiscigralhr, the
center of speciation and generation in this
lineage has been southern South America.
Besides being separable on external morph-

ological and zoogeographical grounds, the
members of the North American lineage,
except for Myiophohus, Cnemotricctis and
NiittoUornis, all have type 3 palatines,
while none of the South American lineage
ha\e them.

I begin the Fluvicolinae with Onycho-
rynchus, a genus that is highly specialized
but apparently is an early offshoot of the
primitive stock. Both sexes of OnycJiorJiyn-
chits have long, elaborate transverse crests,
red or yellow tipped with iridescent blue or
violet, and long, broad, flattened bills with
heavv rictal bristles reaching the tip of the
bill. 'Ames (1971: 160) placed Ouycho-
rhynchus in the Myiohius group with
Terenotriccus and PyrrJiomyias, but Warter
( 1965: 37) made it a monotypic tribe in his
Platyrinchinae, and said it most resembled
Platyrinchus and Tohnoniyias. OnycJio-
rJuiiicJ}Us shares the type 3 nasal septum
with Platyrinchus, and this type is also sug-
gested in Tolmomyias. However, Omjcho-
rhynchus lacks the t)'pe 4 interorbital
septimi, which characterizes the Elaeniinae
in general and is found in Platyrinchus and
allies. On external morphology, Hellmayr
placed Onychorhynchus near Myiohius,
and they certainly share the most extreme
development of the rictal bristles, which ex-
tend beyond the tips of the broad flat bills.
The syringeal e\idence supports this asso-
ciation of the two genera. I keep Onyclio-
rhynchus in the Fluvicolinae, even though
it is the only genus in the subfamily, except
for the aberrant Muscigralla, that does not
have a type 6 nasal septum.

Wetmore (1972: 532) notes that the eggs
of OnycJiorJiynchus and Lophotriccus re-
semble each other in having a "bright to
dark reddish gray ground color, thickly
marked with carmine in scrawling lines," a
tN'pe not found elsewhere among the Tyran-
nidae. This suggested to him that a possible
relationship might exist between the
genera. Eisenmann (in litt.) notes that
Lophotriccus displays its crest in hostile
situations in the same manner as Onycho-
rhynchus, by spreading it laterally, opening

160 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

its beak, and rotating the head through
180°. Although these shared cliaracters are
suggestive, they are not supported by any
anatomical or external morphological char-
acters. The two genera do not share any of
Waiter's cranial characters, and Ames has
them in different syringeal groups — Ony-
chorhynchus in his Myiohius group without
intrinsic muscles, and Lophotriccus in his
Colopteryx group with intrinsic muscles.
Externally, OnycJiorhyncJius is much larger,
with wing length about 60 per cent longer
than that of Lophotriccus, and has a pro-
portionately longer bill, about 32 per cent
of wing length compared to alwut 25 per
cent in Lophotriccus. The tarsus of Ony-
chorJujncJnis is comparati\'ely much shorter
than that of Lophotriccus, and the most
striking difference in proportions is seen in
the ratio culmen tarsus, which is about 155
per cent in Onychorhynchus and about 80
per cent in Lophotriccus. Although the
long crests are used in the same way, they
seem to ha\'e evolved differently. That of
Onychorhynchus appears to ha\'e devel-
oped from a bright orange or yellow crown
of the type found in Tyrannus, being com-
posed of bright feathers greatly lengthened
and tipped with iridescent blue. The crest
of Lophotriccus, on the other hand, is made
up of lengthened ordinary crown feathers,
edged with rusty or gray. Both genera
make pensile nests, but this is true of all
their potential relatives, Myiohius, Rhyn-
chocyclus and Ames' Colopteryx group. Al-
though it is unusual to find simultaneous
convergence in two such unrelated char-
acters as egg color and crest display, I do
not believe that OnycJiorJiynclius is closely
related to Lophotriccus.

The genera Myiohius and Terenotriccus
form a natural group because they and
Onyc]ior])y)ichus are the only genera of
Fluvicolines to make a pendent, purse-
shaped nest, similar to that found in the
Todirostrum group of the Elaeniinae. Myio-
triccus closely resembles Myiohius in form
and plumage pattern, and I include it here
even though its nest is unrecorded. I do not

merge the two because Myiotriccus is more
brightly colored and lacks the heavy rictal
bristles, extending beyond the bill tip, that
are characteristic of Myiohius. This group
of genera is related to Onychorhynchus on
the one hand and to Pyrrhomyias on the
other by their peculiar syringes, in which
there are no intrinsic muscles.

The remaining genera of Warter's "Con-
topini," formerly in the Myiarchinae of
llellmayr, are an unusually difficult group
to order lineally. The majority seem to have
characters relating them to two or more
of the others, and there are a few about
which little is known that must be inserted
next to their apparent nearest relatives,
thereby interrupting an otherwise orderly
progression. Webster ( 1968 ) , in his review
of the genus Mitrephanes, suggested the
following sequence for the genera that he
considered closely related (I invert his
sequence to make it comparable with
mine): Enipidonax, Xenotriccus, Fyrrho-
nujias, Mitrephanes, Contopus, Nuttallornis.
I agree with this sequence, except that I
would place Empidonax at the other end,
next to Contopus and Nuttallornis. Not
only are these genera close in appearance,
but an intergeneric hybrid between Conto-
pus and Enipidonax has been recorded by
Short and Burleigh (1965). My sequence,
including those genera not considered by
Webster, is Myiophohus, Aphanotriccus
(including Praedo), Xenotriccus (including
Aechmolophus), Pyrrhomyias, Mitrepharws,
Contopus (including Nuttallornis and Bla-
cicus).. Enipidonax, Nesotriccus, Cnenio-
triccus, Sayornis and Pyrocephalus.

Myiophohus is a genus that is superfi-
cially like Myiohius, but probably not
closely related. The two are similar in pro-
portions and general appearance, but Myio-
pholnis lacks the bright crest and rump
characteristic of Myiohius, and makes a
cup-shaped rather than a pensile nest.
Myiopholnis is not placed by Ames in any
of his syringeal groups, but Myiohius is
part of the group lacking intrinsic muscles.
It is risky to generalize about Myiophohus,

Classification of Tyrant Flycatchers • Traylor

i<u

because the only species for which the
syrinx and cranium were available, and
whose nest and habits have been recorded,
is fasciatus; the least typical ecologically.
Of the nine species of Myiophobus, all but
fasciatus are found in the subtropical zone
of the Andes, where several ha\'e notably
restricted and fragmented ranges. On the
other hand, fasciatus, is found throughout
the drier lowlands of South America, even
reaching Costa Rica.

Aphanotriccus (including Praedo) and
Xenotriccus (including Aechmoloplius) are
each composed of two relict species with
restricted ranges in Central America and
adjoining Colombia. Both were included
in Ames' NuttaUornis group of genera on
syringeal characters. The species of
Aphanotriccus show the same type of color
variation, in which cinnamon and oli\'e re-
place each other, as is found in Pyrrho-
mijias and Mitrephanes. One species of
Xenotriccus is dull colored like Contopus.
Their relict distribution suggests they may
be remnants of an earlier stock from which
the currently successful Central and North
American genera Contopus, Empidonax
and Sayornis were derived.

Pyrrhomyias cinnamomea is a difficult
genus and species to place because it com-
bines the peculiar syrinx of Ames' Myiohius
group, which lacks intrinsic muscles, with
the form and coloration of Mitrephanes. In
the Myiohius group it would be the only
species that makes a cup-shaped instead of
a pendent nest and has a pointed instead of
a much rounded wing, with 10th primary
equal to the 5th or 6th in length, rather
than shorter than the first. In its pointed
wdng, and even more, in its comparatively
very short tarsus, it agrees with Mitre-
phanes and Contopus. I can find nothing
recorded on the habits of ci7inamomea, an
Andean subtropical species from \^enezuela
to Bolivia, but Mitrephanes phaeocercus is
reported by ^^^ebster ( 1968 ) and Eisen-
mann ( in litt. ) to be a miniature Contopus
in habits.

Contopus (including NuttaUornis and

Blacicus), Empidonax, Pyrocephahis and
Sayornis comprise what I have been calling
the Central and North American genera.
All but Pyrocephahis were included in
Ames' NuttaUornis group on the basis of
syringeal characters. Warter (1965: .36)
also removed Sayornis and Pyrocephahis
from Hellmayr's Fluvicolinae to the vicinity
of Contopus and Empidonax on cranial
characters. The removal of Sayornis from
the restricted Fluvicolinae on zoogeograph-
ical grounds was suggested by Ihering in
1904 (p. 320), but his idea was ignored by
Berlepsch and Hellmayr. Actually, placing
Sayornis near Contopus and Empidonax is
a familiar an-angement to Nortli American
ornithologists, because the American Or-
nithologists' Union Checklist (1957) and
Mayr and Short (1970) place it there.

Sayornis and Empidonax are the most
truly North American fhcatchers. All three
species of Sayornis breed in North America,
and one of them, nigricans, has recently
invaded South America along the Andes,
where it differs only subspecifically. There
are 10 species of Empidonax breeding in
North America, six in Central America and
only two in South America; possibly the
latter represent an invasion from the north,
but the evidence is not so clear as in the
case of Sayornis nigricans. The species of
Contopus are more evenly distributed geo-
grapliically, but the genus is closely related
to Empidonax, as witness an intergeneric
hybrid reported by Short and Burleigh
(1965). Considering present distribution,
Contopus may well ha\'e been the primitive
stock from which the others were deri\ed.
Pyrocephahis has a wide but disjunct range
from southern North America through
drier tropical South America, and even in
the Galapagos; its geographical origins are
not clear, but it bears a close resemblance
in general form and in cranial characters
to Sayornis.

Cnemotriccus fuscatus is found through-
out the tropical lowlands of South America.
In plumage, fuscatus is \ex\ close to Em-
pidonax, but it has a white superciliary in-

162 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

stead of the white eye-ring characteristic
of tliat genus, and has also a blackish
mandil:)le and a rounded rather than square
tail. Hellmayr (1927: 225) included
poecihirus in Cncmotriccus but Zinimer has
shown (1937b: 26) tliat this species is a hen-
colored Knipolcfius. Proportions are so
constant within the medium-sized Fluvi-
colinae that plumage pattern and color and
bill form are frequently the best guides to
relationships.

Nesotriccus rich^uaiji.. tlie endemic genus
and species of Cocos Island, seems to fit
best in the vicinity of Cnemotriccus and
Empidonax, despite cranial evidence to the
contrary. Nesotriccus has always been as-
sumed to be an island representative of
Myiorchtis, but the reasons are more infer-
ential than logical. The Geospizinae of the
Galapagos have an obvious representative
in Pinaroloxia of Cocos Island, and when an
endemic flycatcher was discovered on
Cocos, it was assumed (Townsend, 1895:
124) to be a representative of Eribates
(= Mijiarchus) magnirostris of the Galapa-
gos. Magnirostris is a typical Mijiarchus in
plumage, and differs only in Jiaving the
proportionately longer bill and tarsi char-
acteristic of island forms. Nesotriccus, how-
ever, is not at all Myiarchine in plumage,
but most nearly resembles Empidonax
euleri or Cnemotriccus ftiscatus in both
adult and juvenal plumages. The bill and
tarsi are even more lengthened proportion-
ately than in Eribates. The nest form would
give the best evidence of affinities, but un-
fortunately the nest of Nesotriccus is un-
known. Swarth (1931: 84), comparing the
behavior of ridguayi and magnirostris
writes, "The supposed resemblance be-
tween these two species is non-existent to
me," and Slud (1967: 286) says Nesotriccus
suggested to him a long-billed, slim Em-
pidonax; it moved about in the foliage in
finch-like hops and spurts, and proved
itself adept at aerial feeding, making
flickering short sallies. Nesotriccus lacks
the type 6 nasal septum characteristic of
the Fluvicolinae, but its other cranial char-

acters are sufficiently mixed that Warter
(1965: 37) considered it intermediate be-
tween Mijiarchus and Todirostrum. I shall
leave Nesotriccus between Cnemotriccus
and Empidonax.

The mergings of genera indicated above
have all been previously suggested. NuttaJ-
lornis has been merged with Contopus by
Phillips, Marshall and Monson (1964: 90)
and Blacicus was placed in Contopus by
Bond (1943: 117). Aechmolophus was
made a synonym of Xenotriccus by Web-
ster (1968: 289). Praedo was made a
synonym of Aphanotriccus by Griscom
(1929: 176) although Wetmore (1952: 487)
continues to recognize the fomier.

The remainder of the Fluvicolinae are
the South American element that histor-
ically made up the restricted subfamily of
that naiue. It is composed of a number of
specialized, terrestrial genera with numer-
ous sympatric species, some more general-
ized, arboreal forms and several distinctive
monotypic genera. Despite the great vari-
ation in size, proportions, plumage and
behavior, the Fluvicolines seem a closely
related group. Ames (1971: 158) includes
a broad spectrum of genera in his Fhwicola
group, which is of more significance than
the fact that many closely related genera
were not part of the group. Warter found
that, with the exception of Muscigralla, all
had a type 6 nasal septum, and types 1 or
2 palatines. Geographically, all belong to
the dry country or temperate zones of
South America, or are specialized for
marshes or riverine habitats. None occur
in the humid forests of Amazonia, although
Muscisaxicola fluviatilis and Ochthoeca
(Ochthornis) Uttoralis are found on beaches
and on edges of Amazonian waterways.

Smith and Vuilleumier ( 1971 ) reviewed
the evolutionary relationships of a number
of the ground tyrants; \^iilleumier used evi-
dence from external morphology and
ecology and Smith from behavior. They
included in their study the following
genera of Hellmayr: Agriornis, Xolmis,
Muscisaxicola, Mijiotheretes, Cnemarchus,

Classification of Tyrant Flycatchers • Trayior IQc

Neoxohnis, OchtJiodiaeta, Ochthoeca and
Muscigralla. In their conclusions they
merged Mijiothretes, Cnemarchiis and
Octhocliaefa witli Xolmis; they also kept
Pyrope in Xohnis as Hellmayr had it, al-
though Zimnier (Ms.) and Meyer de
Schauensee (1966: 335) recognized it as a
distinct genus. Muscigralla was made a
subgenus of Muscisaxicola, and the species
mtirina was transferred from Xolmis to
Agriornis. Their sequence of genera was
Ochihoeca, Xolmis, Neoxohnis, Agriornis
and Muscisaxicola, since they considered
the arboreal or bush haunting genera more
primitive or generalized than the puna or
steppe inhabiting terrestrial forms.

I agree with Smith and \^uilleumier's ex-
position of the relationships among these
"ground tyrants," but I feel they have gone
too far in lumping some of the genera. All
the species included in Hellmayr's Mijio-
theretes, CnemarcJms and Ochthodiaeta are
characterized by brown, buff or rusty
plumage, and are found only in brush or
the edge of forest in the high Andes, from
Venezuela to northwestern Argentina. The
species of Hellmayr's Xolmis, on the other
hand, are with one exception patterned in
black, gray or white and are found in
tropical to temperate lowlands from eastern
Brazil and Bolivia south to Patagonia. I
feel that the dichotomy between these two
groups is so marked that, even though they
may have had a common ancestor, their
present relationship is best shown by the
recognition of two genera — Mijiotheretes
(including Cnemarchiis and Ochthodiaeta)
and Xolmis.

The one exception to the black, gray and
white plumage pattern in Xolmis is ruhetra,
which is brown above with a russet crown,
and whitish below, streaked dark brown on
throat and breast, and with a russet wash
on the flanks. Vuilleumier (in Smith and
Vuilleumier 1971: 197) says, "In color, pat-
tern and habits, X. ruhetra is intermediate
between the other species of Xolmis and
Neoxohnis rujiventer, and may represent an

evolutionarv transitional

stage

between

arboreal and terrestrial tyrants." Neoxolmis
is almost wholly terrestrial in habits, only
occasionally perching on low bushes, and
it was for this reason, along with the pro-
portionately longer tarsus associated with
terrestrial habits, that Vuilleumier recog-
nized it. Ruhetra is the most terrestrial of
the Xolmis species, with a tarsus propor-
tionately as long as in riifiventris, and its
plumage is not far from the female plumage
of riifiventris. Ruhetra and rufiventris have
allopatric breeding ranges, the former in
northern Patagonia from Mendoza to
Chubut, the latter in southern Patagonia
and Tierra del Fuego. I believe that trans-
ferring ruhetra from Xolmis to Neoxolmis
best shows the relationships of these forms.
In the remainder of the genera treated, I
accept Smith and Vuilleumier's revision
with a single exception, the inclusion of
Muscigralla in Muscisaxicola. As discussed
in more detail below, I find the former so
distinct morphologically that I cannot con-
sider it closely related to any other genus.
I have also accepted Vuilleumier's (p. 192)
tentative suggestions that the monotypic
genera Ochthornis and Colorhamphus be
merged in Ochthoeca. Ochthornis littoralis
is a riverine species foimd throughout
Amazonia, while the species of Ochthoeca
are found in temperate Andean forest or
scrub. Littoralis is dull brown throughout,
but with the white superciliaries of Och-
thoeca; in proportions it has a longer bill
and shorter tarsus than the Andean forms.
In plumage and proportions it bears the
same relation to Ochthoeca that Muscisaxi-
cola fluviatilis (also a riverine form of
Amazonia) bears to the remaining Andean
species of its genus, and I place Ochthornis
in Ochthoeca. Colorhamphus parvirostris
has been kept in the Serpophaginae (now
Elaeniinae), but it also appears to be a
dulled Ochthoeca. Cranial characters,
which might give more positive evidence,
are lacking. In size and proportions parvi-
rostris is identical with Ochthoeca rufi-
pectoralis; however, in plumage it is closest
to oeminthoides, of which it appears to be a

164 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

geographical representative. Oenanthoides
breeds south throiigli the Andes to the
mountains of nortlivvestern Argentina, while
parvirostris replaces it in Patagonia and
Fuegia. Vuilleumier (loc. cit.) also sug-
gested that Tiimbezia be included in Och-
thoeca. The former has a more slender bill
and more pointed wing than the Ochthoe-
cas, and inhabits the arid coastal zone of
Peru along with the peculiar Musciiiralla.
Although these are not trenchant charac-
ters, Tumhezia does not suggest Ochthoeca
to me, and I keep it monotypic.

Lessonia appears to be a link, pheno-
typically at least, between the terrestrial
Muscisaxicola and the more arboreal
Knipolcii^us. In form the Lessonia species
are similar to Muscisaxicola, with long tarsi
and pointed wings; they are completely
terrestrial, and have a lengthened hind
claw like many of the pipits (Anthiis). On
the other hand, they are markedly sexually
dimorphic, the males are mostly black and
the females brownish like the majority of
Knipole^tis.

In Knipolegus I include Phaeotriccus and
Entotriccus, two genera showing the same
type of sexual dimorphism, blue-black bill
and form as Kiiipolegus, but separated
from the latter by the narrow, blade-shaped
outer primaries. In PJiaeotricciis the three
outer primaries are narrowed, as are all in
Entotriccus, but aberrant primaries vary
intragenerically so often in the flycatchers,
that by themselves they serve only as spe-
cific characters. Short (1975: 269) has also
suggested merging the three.

Hymenops {Lichcnops in Hellmayr) is
intermediate between Knipole<ius and the
palustrine Fhwicola and Arundinicola. In
plumage it is similar to Knipolegus, having
a black male and a brown female with
heavily streaked white underparts. How-
ever, the male has a bright vellow bill and
yellow caruncles around the eye, and both
sexes have a short, rounded wing, with the
7th secondary almost reaching the wing tip.
Hymenops is palustrine, as are Fhwicola
and Arundinicola. The latter two, although

differing in proportions and wing form,
are so alike in their black and white or
brown and white plumage, palustrine
habitat and nest form, that I feel they must
be united under the oldest name, Fhwicola.

Following Fhwicola is a group of four
fancy-tailed tyrants that Warter (1965:
130) thinks should be treated as a distinct
tribe, Alectrurini, within the Fluvicolines.
At present they are kept in four monotypic
genera, Colonia, Guhernetes, Alectrurus
and Yetapa. Colonia, with its black and I
white plumage, seems to link this group
with Fhwicola, although Ames said the
syrinx of Colonia was unlike Fhwicola and
more nearly like Ochthoeca. Colonia differs
from the other tliiee species in having the
central instead of the outer rectrices devel-
oped into ornamental plumes, and in being
a short-billed, short-legged, arboreal, hole-
nester, rather than a long-billed, long-
legged, terrestrial, ground-nester. Guher-
netes, Alectrurus and Yetapa have always
been considered closely related. The last '
two should be merged as suggested by
Short (1975: 269); the only differences are
in the shape of the ornamental rectrices,
and the fact that the male of A. tricolor
loses the feathers of the face during the
lireeding season. Guhernetes is much
larger, lacks the black and white plumage
found in the males of Alectrurus, and
has a forked tail like Muscivora tyran-
nus rather than ornamental outer rectrices.
I keep it as a distinct genus. While I accept
the near relationship of Colonia, Guher-
netes and Alectrurus as shown by the
cranial characters, I do not consider them
more distinct than the other groups of
Fluvicoline genera.

There remain in the Fluvicolinae six
monotypic genera that are kept together
not because they are necessarily related to
each other, but because none of them seems
to have any demonstrable close relative.
They are: Satrapa, Tumhezia, Muscigralla,
Hirundinea, Machetornis and Muscipipra.
The first two may be related. Both have
dark, uniform upperparts, olive green in

Classification of Tyrant Flycatchers • Traijlor 165

Satrapa and brown in Tnmbezia, and both
have a yellow forehead, siiperciliaries and
underparts. Satrapa, however, is a larger
bird with proportionately much shorter
tarsus — 20.5 per cent of wing length com-
pared to 29 per cent. Tumbezia occurs on
the arid Peruvian coast and Satrapa in the
dry country east of the Andes and south of
Amazonia. They may, as noted before, turn
out to be related to Ochfhoeca.

Smith and Vuilleumier ( 1971 ) made
Mtiscigralla a subgenus of Mttscisaxicola.
However, Muscigralla brevicauda is unlike
any Muscisaxicola in proportions and plum-
age pattern, and has tarsal scutellation un-
like any flycatcher. It has rounded wings
rather than the very pointed wings of
Muscisaxicola, a proportionately short tail,
and much longer culmen and tarsus. Unlike
the species of Muscisaxicola, which have
uniform dull brownish backs and blackish
tails and tail coverts, brevicauda has pale
buff tipping on the rump feathers, bright
chestnut uppertail coverts, and a blackish
tail tipped with pale buff. It also has a
concealed yellow crest of a type not found
in Muscisaxicola. The most distinctive dif-
ferences, however, are found in the tarsi.
All Muscisaxicola have typically exaspidean
tarsi, with at most a few scutes on the
proximal half of the plantar surface; there
is a deep groove on the inner surface where
the scutes meet, but none on the outer sur-
face, and the scutes are smooth. Musci-
gralla has a classic holaspidean tarsus, with
a deep groove on both inner and outer sur-
faces, and the distal edges of the scutes are
raised, giving a roughened serrate feel.
Furthermore, this scutellation extends for
about 10 mm above the tibio-tarsal joint
as in the shorebirds, a condition found in
no other flycatcher. Ames (1971: 160) be-
lieved its syrinx was most nearly related to
Nuttallornis and not the Fluvicolinae. War-
ter ( 1965: 36) found it to be a good Tyran-
nid, but (p. 31) listed it as the only Fluvi-
coline without a type 6 nasal septum. Its
geographic range, the arid coast from Ecua-
dor to southern Peru, is typical of species

in several groups. Despite the fact that
Smith and Vuilleumier (1971: 201, 253)
and Fitzpatrick (in lift.) consider its dis-
plays and vocalizations closely related to
those of Muscisaxicola, I cannot see that
Muscigralla has any close relatives; morph-
ologically and anatomically it is not typical
of the Fluvicolines. On the other hand, it is
equally out of place in any other group, and
considering its generally brownish color,
terrestrial habits and restricted non-forest
range, it was probably an early offshoot of
the Fluvicoline stock, and I shall keep it at
the end of that group with other aberrant
genera.

Hirundinea is a monotypic genus placed
by Berlepsch ( 1907 ) without comment be-
tween Myiobius and Onychrorhynchus at
the end of the Myiarchinae. I cannot see
any relationship with that group. Hirun-
dinea ferruginea is a pointed winged, aerial
feeder with proportionately the shortest
tarsus of any flycatcher, 12 per cent of wing
length; the bill is long but the rictus only
moderately bristled. It occurs in woodland
and campos north and south of Amazonia,
extending to northern Argentina and tem-
perate Bolivia. It nests primarily on cliff
faces, but in some areas it is virtually a
commensal of man, placing its nest in
niches of buildings or old oven bird nests,
and it will sometimes wall off its nest with
pebbles (Euler, 1900: 48). Onychrorhyn-
chus and Myiobius, on the other hand, are
round winged, forest haunting birds, with
the heaviest rictal bristles of any of the fly-
catchers, and make long, pendent purse-
shaped nests. In proportions, Hirundinea
is more like the Tyrannine flycatchers with
its pointed wing and short tarsi, but it is
unlike any of that group in plumage and
nesting habits. In distribution it is typical
of the Fluvicolines, and I place it with the
other peculiar monotypic genera of that
group. Unfortunately, no one has had
anatomical material available for study.

Machetornis rixosus is a rounded winged,
long-legged, terrestrial species that in
plumage pattern and in temperment is re-

166 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

markably like the tropical kingbirds of the
genus Tyrannus. It would be tempting to
consider it a member of the latter group,
secondarily adapted for terrestrial life, were
it not for the fact that its cranium is
typically Fluvicoline, and it does not share
a single cranial character with the Tyran-
nines. Ames (1971: 147) says that the Mm.
sternotracheales of Machctornis strongly
resemble those of Xohnis^ but that it lacks
intrinsic muscles. He considered that if the
relationship to the cither "ground tyrants"
was real, Machetornls must have lost its
intrinsic muscles. The last of the aberrant
Fluvicolines, Miiscipipra vetula, is again
much like the Tyrannine kingbirds, both in
plumage and proportions. In absolute size
and proportions it is very close to Tyrannus
scwana (Miiscivora tyrannus), but it is dull
gray rather than black and white, lacks any
bright crest, and the fork of the tail is less
deep. On the other hand, it shares the same
proportions with Colonia among the Fluvi-
coHnes, and on the basis of syringeal char-
acters, Ames (1971: 158) placed it in his
Fhwicola group. Warter had no specimen
and I can find nothing on its habits. Its
restricted range in southeastern Brazil and
adjoining Paraguay and Misiones, Argen-
tina, tells us little. Mu^cipipra may well
prove to be a Tyrannine, but in the absence
of more concrete evidence, I leaxe it in the
Fluvicolines.

TYRANNINAE

The remaining subfamily, Tyranninae,
consists of Hellmayr's old Tyranninae plus
Myiarchus and allied genera. WHiile it is
difficult to define the subfaiuily as ad-
vanced or specialized, it is certainly suc-
cessful and is the one that appears to have
undergone the most recent radiation. Al-
ni(xst all the genera and many of the species
are found throughout Central and South
America, and the majority are aggressive
and doiuinant wherever found. The ex-
treme uniformity of the crania is exidence
of recent radiation; only two types of nasal
septa occur, both without basal plates, and

interorbital septa, palatines and cranial
types are identical throughout.

Warter recognized Attilinae as a sub-
family (1965: 95, 140) because of its
peculiar cranial characters; it included
Attila, and presumably Pseudattila and
Casiornis, which Warter was unable to
examine. However, as noted before, ex-
ternal morphology and syringeal characters
strongly suggest that Attila (including
Pseudattila), Casiornis, Laniocera and Rhy-
tipterna are a closely related group of
genera belonging with Myiarchus. Rhytip-
terna and Myiarchus in turn are clearly part
of the subfamily Tyranninae, as evidenced
by cranial characters. Despite the contrary
cranial evidence separating Attila from the
rest of the Myiarchus group, I beUeve the
weight of evidence is in favor of keeping
them all together.

I begin the Tvranninae with the Miliar-
chus group. Syristes, which has previously
been placed near Tyrannus by Hellmayr
(1927: 119), but in the Cotingidae by
Ridgway (1907: <S15), also belongs here.
It has a holaspidean tarsus similar to
Casiornis or Rhytipterna, and the cranial
characters of Syristes and Rhytipterna are
identical. Although the plumage is more
like that of Tyrannus, the rounded wing,
holaspidean tarsus and type 2 nasal septum
place it with Rhytipterna and near Myiar-
chus. This is essentially the sequence of
Wetmore (1972: 433), who put it between
Myiarchus and Attila. Hylonax and Eribates
are included in Myiarchus. Deltarhynchus
flamnuilatus, which is pure Myiarchus in
plumage, is separated from the latter by its
short, triangular bill. Unfortunately noth-
ing has been published on the anatomy or
behavior of flamnuilatus. If it is found to
be a hole nester, using a few scraps of snake
skin for decoration, it should certainly be
luerged in Myiarchus.

The remainder of the genera are those
included by Hellmayr in his Tyranninae.
Although they are related to the Myiarchus
group through identical interorbital septa,
palatines and crania, they differ, with the

Classification of Tyrant Flycatchers • Trayior

if;7

exception of Legatiis, in having a type 1
instead of a type 2 nasal septum, and the
majority belong to the Tyrannus syringeal
group rather than the MyiarcJuis group.

Meise (1949) reviewed Hellniayr's Ty-
ranninae, and separated the genera into
three groups on the basis of nest type. They
were: 1) those with open, bowl-shaped
nests — Tyranmis (including Tyrannopsis
and TolmarcJjus), Muscivora, Empidono-
mtis, Legatus, Conopias (including Cory-
photriccus) and MegaHiynchus: 2) hole
nesters — Myiodynastes\ and 3) those with
globular nests — Myiozetetes and Pitangtis.
Although Meise's classification is a useful
approach, increasing knowledge has made
some of his conclusions questionable. Cono-
pias parva turns out to nest in holes and
nested at least once in an abandoned ca-
cique nest ( Haverschmidt 1957: 240; 1973:
207). Within the genus Pitangus, siilphur-
atus makes a globular nest and lictor
usually a cup-shaped nest. The nest de-
scribed for Conopias- inornata by Cherrie
(1916: 238), a small cup, saddled on a
branch and neatly covered with lichens, is
totally imlike the loose, stick nests in forks
or crotches characteristic of other cup-nest
builders in this subfamily. Finally Legatus
usurps the domed nests of many species
from different families, annoying the
owners until thev abandon the completed
nest (Skutch, 1960: 451; Haverschmidt
1968: 305). Although nest type is an im-
portant character, it cannot be the primary
basis for a classification.

There are five currently recognized
genera in which the same distinctive plum-
age pattern occurs in at least one species.
These are Pitangus, Myiozetetes, Conopias,
Myiodynastes and Megarhynchiis. The
plumage characteristic of this group of
genera has the crown blackish with a con-
cealed yellow or orange crest; white fore-
head and superciliaries, with the latter usu-
ally extending around to meet on the nape;
broad black band tluough the eye; upper-
parts olive to brown, in contrast with the
dark crown; wings and tail brown, the

feathers edged with the color of the back or
with chestnut; throat white or yellow;
breast, belly and crissum bright yellow.
This pattern is found inter alia in Myio-
dynastes hemichrysus, Megarhynchiis pi-
tangua, Conopias parvus, Myiozetetes
cayannensis and similis, and Pitangus sul-
pha ratus and lictor.

In Table II the species of these genera are
listed with some of the character states used
to separate them. The table shows the spe-
cies in each genus are for the most part
more closely related to each other than they
are to any outside species; what is not clear
is how many genera should be recognized.
Myiodynastes is separated from all others
except Megarhynchiis by large size, long
stout bill and short tarsus (except for
hairdii), and from all except Conopias par-
vus by its hole-nesting habits. In external
form Megarhynchiis is merely an exag-
gerated Myiodynastes with a long, deep,
decurved bill, but it does make an exposed,
cup-shaped nest high up in trees, a typical
Tyrannus-tyipe nest, in contrast to the hole
nesters. Fitzpatrick ( in litt. ) says Mega-
rhynchiis is much closer to Pitangus in be-
ha\ior, since it feeds in the open, rather
than within the foliage as does Myio-
dynastes. The three typical Myiozetetes,
cayannensis, granadensis and siiniUs, are
well defined by short bill, globular nest and
syrinx unlike any other; the nest of lutei-
ventris [placed in Tyrannopsis by Zimmer
(Ms.), and Meyer de Schauensee (1966)]
is imknown. However, they share the short
bill with Conopias inornatiis (nest cup-
shaped), and the globular nest with Pitan-
gus sulpliuratus (large and long-billed).
Conopias is the least known genus as far as
behavioral or anatomical characters go.
C. parva is a hole nester; if this holds for
the other two "typical" species, then Cono-
pias is probably more closely related to
Myiodynastes than to Myiozetetes. C. in-
ornatus has the size and proportions of
Myiozetetes, but lacks a brightly colored
crest, as do Conopias trivirgata and chin-
choneti for the most part, and has yellow

168 Bulletin Museiiiii of Comparative Zoology, Vol. 148, No. 4

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Classification of Tyrant Flycatchers • Traylor 1C3

rather than buff inner edges to the pri-
maries. Based on one record (Cherrie,
1916: 238), inornatus liuilds a neat shallow
cup nest, covered with lichens, and saddled
on a large limb. If true, this sets inornatus
apart from either Conopias or Myiozetetes.
For the moment I leave inornatus in Cono-
pias, although Zimmer (Ms.) and Meyer
de Schauensee (1966) place it in Myio-
zetetes. A final problem in this group is the
status of the two Pitangus species, sulphur-
atus and lictor. The two are virtually
identical in plumage and proportions, and
have more slender bills than other tyran-
nines. However, siilphiirafiis builds a globu-
lar nest, and lictor an open cup ( Haver-
schmidt, 1957: 240; Smith, 1962: 111); the
eggs of lictor are a much broader oval than
those of sulphuratus (Meise, 1968: 78);
sulphuratus has a typical tyrannine syrinx,
but according to Ames (1971: 159) "Pitan-
gus lictor lacks most of the features of the
[Tyrannus] group and, in view of the
structural homogeneity among those listed
above, it is difficult to believe lictor and
sulphuratus are more closely related to each
other than the latter is to Tyrannus"; and
Warter (1965: 36) says the crania of lictor
and sulphuratus differ more than any other
congeners . If lictor and sulphuratus are not
congeneric, there is no logical genus in
which to place the foiTner. With its slender
bill, it is the antithesis of the other cup-
nest maker Megarhynchus, and on syringeal
characters it differs as much from Myio-
dynastes, Megarhynchus and Conopias as it
does from P. sulphuratus. Myiozetetes also
differs from the tyrannines on syringeal
characters, but not in the same way as
lictor, and the nest of the latter is unlike the
globular nest of Myiozetetes. Wetmore
(1972: 422) says lictor should probably be
placed in a separate genus, but considering
my imperfect knowledge of this and related
groups, I cannot see creating a new genus
at this time. I shall leave lictor in Pitangus,
Init only for lack of a better place to put it.
I recognize the above five genera, not
because I consider them satisfactory, but

because at the moment I have nothing
better to suggest. A case can be made for
uniting Myiozetetes with Conopias, or
Pitangus with Myiozetetes, but as noted in
the discussion above, there are good
reasons for not doing so. Until more is
known about the behavior and nesting
habits of some of the more obscure species,
I feel more comfortable leaving them as
they are. They are listed in the order
Pitangus, Megarhynchus, Myiozetetes,
Conopias and Alyiodynastes.

Legatus leucophaius is almost identical
in both Juvenal and adult plumages with
Empidonormis varius: the only noticeable
difference is the pale buff edge on the inner
webs of the primaries in leucophaius con-
trasted with the yellow edge in varius. In
size, however, leucophaius is smaller, with
a proportionately shorter, broader bill,
shorter tail but longer tarsus. They differ
markedly in nesting habits, leucophaius
usurping the domed nests of other spe-
cies, while varius builds an open cup. War-
ter (1965: 32) found Legatus had a type 2
nasal septum like Myiarclius, rather than
the type 1 found in the rest of the restricted
tyrannines, and Ames (1971: 159) reported
the syrinx to be unlike the rest of the Ty-
rannus group. Considering these differ-
ences, the similarity in plumage is probably
due to convergence.

Empidonoinus varius and aurantioatro-
cristatus are closely related to Tyrannus.
They are an obvious species pair, being
almost identical in measurements, and each
having a long blackish crest with bright
yellow vertex; varius, however, is streaked
above and below, while aurantioatrocrista-
tus is plain brown above and gray below.
Meise (1949: 71) considered Empidononuis
separable from Tyrannus by its short bill
and long tail. The former does have a short
bill compared to the Tropical Kingbird, T.
melancholicus, but no shorter than the
North American Kingbird, T. tyrannus, the
bill/wing ratios being 17-18 per cent.
Similarly, the tails of Enipidonomus species
are proiDortionately the same as that of T.

170 Bulletin Museum of Conipdrative Zoology, Vol. 148, No. 4

melancholicus, 81-S3 per cent of wing
lengtli. Syrinx, cranium, notching of pri-
maries, and nest form are the same in l)otli
genera. Furtlier evidence of the close re-
lationship of Empidonomus and Tyranmi.s
is tlie fact, demonstrated by Meise (1949:
61) that Laphijctes apoUtes Cabanis and
Heine is a hybrid between E. varius and T.
melancholicus. The one character I con-
sider sufficient to maintain their separation
is the much more slender build of Empido-
nomus. When specimens of the two genera,
prepared by the same collector, are com-
pared, those of Empidonomus are invari-
ably more slender and cigar-shaped, while
those of Ti/rannus are broader and more
chesty. While such a difference is not
precise, it must reflect some anatomical
and probably behavioral difference, and
for the moment I recognize the two genera.
The streaked ]:)lumage of varius would also
be out of place in Tijrannus.

Tyrannopsis is a peculiar monotypic
genus that is apparently most closely re-
lated to Tijrannus., although the single spe-
cies sidphurea was for a long time carried
in Myiozetetes. It differs from Myiozetctes
in being larger, with a proportionately
shorter tail and longer culmen, in having a
syrinx belonging to the Tyrannus group,
while that of Myiozetetes is unlike any
other, and in building a T/yra/i /H/s-like cup-
shaped nest instead of a globular one. In
plumage, sulphuraea is a saturated version
of Tyrannus melancholicus with a dark
gray head and orange crest, dark olive
Ixick, gray throat with a white center, and
yellow breast and belly. WHiere the gray
of the sides of the throat bleeds into the
yellow breast, there is oli\'e streaking, a
character not found in any of the kingbirds.
In size and proportions Tyrannopsis fits
within the overall limits of Tyrannus, but is
not close to any one species, having the
short bill of the North American tyrannus
and verticalis, and the long tarsus of the
insular caudifasciatus. The male lacks the
notched primaries present in all continental
Tiirannus. The range of sidj)Jiurca is exten-

sive in the tropical lowlands east of the
Andes, south to southern Amazonia and
east to Maranhao, Brasil, but its distribu-
tion is restricted to areas where the Mauri-
tia palm, in which it nests, is found. Meise
(1949: 71) placed sulphurea in Tyrannus,
but Smith (1966) in his review of that
genus did not mention it as a possible Ty-
rannus. To further complicate its possible
relationships, Zimmer ( Ms. ) followed by
Meyer de Schauensee ( 1966: 346) added to
Tyrannopsis Myiozetetes hiteive^itris, a spe-
cies that by no stretch of the imagination
could be considered a Tyrannus. Luteiven-
tris is a miniature of sulpJuirea in colora-
tion, even smaller than the smallest Myio-
zetetes, and it is identical with the latter in
proportions; its nest is not known. I feel,
as did Blake (1961), that luteiventris should
remain in Myiozetetes. I also recognize
Tyrannopsis as a genus, at least until it is
better known behaviorally.

The last genus in the family is Tyrannus.
It is the most widespread geographically,
both as a genus and in the individual
ranges of some species, and the species are
usually aggressive and dominant wherever
they occur. Considering the uniformity of
the species, both morphologically and be-
haviorally, they probably represent the
most recent radiation in the family.

Smith (1966) comprehensively reviewed
communications and relationships within
the genus Tyrannus. He merged with Ty-
rannus the monotypic Tolmarchus of the
West Indies, and Muscivora with its two
fork-tailed species, tyrannus and forficata.
The merging of Tobnarchus with Tyrannus
had been previously recommended by
Meise (1949: 73) and Bond (1958); it is
supported by syringeal and cranial evi-
dence. Smith's merging of Muscivora has
had a mixed reception, being followed by
Lanyon (1967a: 606) and by Mayr and
Short (1970: 59), but opposed by Howell
(1972: 325). The union of Tyrannus with
Muscivora recjuires that Muscivora tyran-
nus \)v called Tyrannus .savana, to avoid
h()mon\ni\ with the North American Ty-

Classificatiox of Tyrant Flycatchers • Traylor

ranntis ti/rannus. Smith's decision was
based on behavioral considerations, but
other evidence is equally strong. Cranial
characters, syrinx, nest form and notched
^ primaries are alike in Muscivora and Tij-
r(ninu.s-. Except for the long, forked tail, the
Muscivora species are almost identical in
size and proportions with the North Ameri-
can Tyronniis tyrannus and verticalis. This
is of particular interest, since Meise (1949:
75) belie\ed .savana and forficata evolved
independently from T. tyrannus and verti-
calis respectively. Further evidence of
close relationship is found in the hybrid be-
tween verticalis and foiiicata described by
Davis and Webster (1970). If savana and
forficata did evolve independently, then
their long forked tails are convergent, and
there is no taxon Muscivora. At present I
do not know of any characters that could
be used to prove or disprove this hypoth-
esis. I do not recognize Muscivora e\en as
a subgenus because of its close relation to
Tyrannus, but I keep savana and forficata
together because of their possible common
origin.

, PHYLOGENY

Any attempt to derive a phylogeny re-
quires some knowledge of the primitive
and derived states of the characters on
which the classification is based. Also, it
must be possible to determine whether the
derived state is monophyletic or whether it
evolved independently two or more times,
and as a corollary, whether a state is really
primitix'c or just a secondary regression
from a derived state. When the characters
used to classify the Tyrannidae are viewed
in this light, there are few that serve for
determining a phylogeny.

There are a number of morphological
characters that occur regularly in the
family, but the characters are so randomh'
distributed they must have evolved or been
lost independently many times. Bright
coronal crests, aberrant primaries, and
hola.spidean, taxaspidean or pycnaspidean
tarsi are found in unrelated groups in all

three subfamilies, and merely represent a
genetic potential within the family for this
type of mutation. Even the pendent, pyri-
form nest, which is an elaborate structure
requiring a special set of behavioral pat-
terns for its construction, must have evolved
at least twice — in the Myiohius group of
the Fluvicolinae and in the tody-tyrants of
the Elaeniinae. As Ames (1971: 150)
noted, "The application of syringeal mor-
phology increases in reliability toward the
generic level," and he made no attempt to
rate his groups of genera as primitive or
advanced. The only characters that seem
amenable to classification as primitive or
derived are the cranial characters of War-
ter.

In classifying the states of the cranial
characters, I have used the e\ddence of
their present distribution within the sub-
families, rather than tr\ ing to determine
which are intrinsically primiti\e or derived.

1 ) Nasal septum - types 1 and 2 are

found in significant proportions in
both Tyranninae and Elaeniinae,
while a type 6 septum is found in
all the Fluvicolinae except the two
aberrant genera Muscig,raUa and
Onychorhynchus. Types 3 and 5
occur once each in the Fkuicolinae
and occasionally in the Elaeniinae.
Type 1 or 2 is almost certainly
primitive, and the others derived,
but only type 6 appears significant
phylogenetically.

2) Interorbital septum - type 1 occurs

regularly in the Fluvicolinae, ex-
clusively in the Tyranninae and
twice in the Elaeniinae. Type 2 is
mostly confined to the Fluvicoli-
nae, but occurs twice in the
Elaeniinae; type 3 is found occa-
sionalK- in both, but type 4 is found
only in the Elaeniinae, where it
occurs in 77 per cent of the studied
genera. E\'idently type 1 is the
primiti\'e type, type 4 is derived
and characterizes the Elaeniinae,
and types 2 and 3 ha\e evolved

172 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

WING LENGTH
80 90

100

110

120

130

140

z
<

■

TYRANNIN

(a)

•

FLUVICOLI

(b)

NAE

•

151

•
•

• •

•

• • •

ELAENIINA
(c)

•

•
•

• •

• ••

Figure 6. Warter's cranial types plotted as a function of wing length for the three subfamilies of Tyrannidae.
In the Tyranninae (a) only type 1 occurs, and it is characteristic of the subfamily. In the Fluvicolinae (b) and
Elaeniinae (c), the cranial types show a close correlation with size, as represented by wing length, and are
without taxonomic significance.

independently in Fluvicolinae and
Elaeniinae.

3) Palatines - type 1 occurs regularly in

all three subfamilies, and is the
only type found in the Tyranninae;
it must be the primitive type. Type
2 is also common in the Fluvicoli-
nae and Elaeniinae, but type 3 is
confined with one exception to the
"Contopine" lineage of the Fluvi-
colinae.

4) Cranium - with the exception of the

Tyranninae, the cranial types are
strongly correlated with size as ex-
pressed by wing length — type 1
being found in the largest birds
and type 3 in the smallest (Fig. 6).
In the Tyranninae only type 1

occurs, regardless of wing length.
Assuming the primitive flycatchers
were small arboreal forms, nearest
the present Elaeniinae, then types
2 and 3 are primitive, and type 1
derived.

Figure 7 is a diagram suggesting the
possible origin of the subfamilies of fly-
catchers, incorporating the primitive and
derived states of the cranial characters. The
putative proto-flycatcher appears within
the circle, with the different subfamilies
leading off from it. The derived cranial
characters defining the subfamilies are
indicated, and in two cases the subfamilies
are further subdivided. Finally, Ames'
seven syringeal groups are indicated, show-
ing the close correlation between the two

Classification of Tyrant Flycatchers • Tray

1 .'""r^

±.'0

AMES TYRANNUS

GROUP
NASAL SEPTUM 1

AMES' MYIARCHUS

GROUP
NASAL SEPTUM 2

AMES f LUVICULA
GROUP
PALATINES 1 2

AMES' NUTTALLORN^

and MYIOBIUS GROUPS

PALATINES 3

FLUVICULINAE
NASAL SEPTUM 6

INTERORBITAL SEPTUM 4

ELAENINAE
AMES' COLOPTERYX and
ELAENtA GROUPS

Figure 7. Postulated origin of the subfamilies of
Tyrannidae, based on Warter's cranial characters.
Ames' groups of genera, based upon syringeal char-
acters, are added to show the close correlation be-
tween these character complexes.

types of anatomical characters. It is tempt-
ing to try to carry out further subdivisions
within the subfamiHes, based on cranial or
other evidence, but it is unprofitable for
two reasons. First, the e\ddence, even
within the cranial characters, becomes con-
flicting, and second, there are too many un-
examined genera whose allocation would
be guesswork.

The circular form of diagram in Figure
7 is used because the three subfamilies of
flycatchers seem to have arisen indepen-
dently rather than sequentially, one from
another. There are no genera in any given
subfamily that seem ancestral to or even
closely related to either of the other sub-
families. Onychorhi/nclius was placed in
my Elaeniinae by Warter, and in the Fluvi-
colinae here, but the difficultv is not that
Oniichorhynchiis is intermediate between
tlie two, but that its high specialization
makes it difficult to place it in either. If
my reconstruction of the proto-flycatcher
characters is correct, then none of the
modern taxa retain the primitive condition.

Elaeniinae — This subfamily is charac-

terized by the type 4 interorbital septum,
which is found in 27 of the 35 genera ex-
amined by Warter. The remaining genera
have variously types 1, 2 or 3, and the evi-
dence suggests these represent a secondary
loss of type 4 rather than the retention of a
primitive state. Seven of these genera —
Myiopagis, Elaenia, Suiriri, Su])leg,atus,
Phaeomyias, Zimmerius and Tyranmdus —
belong to the old restricted subfamily
Elaeniinae, and all but Sublegatm and
Zimmerius are part of Ames' Eloenia syrin-
geal group. However, their relationships
are otherwise divided among groups still
retaining the type 4 interorbital septum.
Elaenia, Suiriri and Sublegatus share a type
5 nasal septum with the nearly related
Anairetes, Serpophoga, Inezia and Meco-
cercuJus. This type of septum is certainly
derived and is found only here, with the
exception of the peculiar Miiscigralla of the
Fluvicolinae. Phaeomyias, on the other
hand, is most closely related to Campto-
stoma, which has a type 4 interorbital sep-
tum. The last genus to lack the type 4
interorbital septum is Polystictus, which is
related to P.seudocolopteryx, which has the
normal type. A second of Ames' syringeal
groups, the Colopteryx group, includes
most of the tody-tyrants, all of which have
the type 4 interorbital septum.

Fluvicolinae — The Fluvicolinae are
characterized by the type 6 nasal septum,
which is found in all genera except the
highly specialized MuscigraUa and Onycho-
rhynchiis. This type of nasal septum has also
evolved independently twice in the Elae-
niinae, in Polystictus and in Zimmerius.
The fact that these two genera have the
Fluvicoline nasal septum and lack the char-
acteristic type 4 interorbital septum of the
Elaeniinae might suggest that one or both
are primitive links between the Elaeniinae
and Fluvicolinae. However, in general con-
formation and appearance they are typi-
cally Elaeniinae and unlike any Fluvicoli-
nae, so I believe the resemblance in cranial
characters is due to convergence. Certainly
it would be difficult to conceive of both

174 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

representing the same primiti\'e stock, since
Poh/sticttis is a l:>rown, streaked grassland
form, and VAmmeriii.s a green, forest group.

Tlie Fluvicolinae are further subdivided
by the possession of tlie derived type 3
palatine in Warter's "Contopine" lineage.
This palatine occurs elsewhere only in
Platyrinchtis of the Elaeniinae, where it is
almost certainly independently derived.
Three of the genera of the "Contopines,"
NutfaUornh; Cnemotriccus and Myiopho-
hus, have a type 2 palatine, but this almost
certainly represents a secondary loss of the
type 3. This is most evident for Niittallornis,
whose congeners in Contopus all have type
3 palatines, and for Cnemotriccus, whose
closest relative is Empidonax with type 3
palatines. Myiophohtis does not have any
single close relative, but the genera closest
to it on other characters, Mitrephanes,
Pyrrhomyias and Myiobius, all have type 3
palatines.

Ames' syringeal groups support this
derivation of the Fhu'icolinae. Two of
them, the Myiobius group and the Nuttal-
lornis group, are part of the "Contopine"
lineage, while the Fhwicola group falls in
the restricted Fluvicolines.

Tyranninae — The Tyranninae are re-
markable in that all genera, with the excep-
tion of Attila, have one of two invariant
sets of cranial characters. All but Attila
have type 1 interorbital septa, palatines and
crania, and these are associated with either
type 1 or 2 nasal septa. The restricted
Tyrannines all have type 1 nasal septa, with
the exception of Legotu.s, and the Myiar-
chine group have type 2, with the exception
of Attila. The latter type is presumably the
derived form, since the type 1 is more com-
mon among the Elaeniinae. Ames' syringeal
groups fit this dichotomy of the Tyran-
ninae. The Myiarchus group includes those
genera with type 2 nasal septa, plus Attila,
and the Tyrannus group includes only
genera with type 1 nasal septa, the
restricted Tyranninae.

The most interesting aspect of the Tyran-
nine cranium is the extreme uniformity, as

if all the character states were inherited as
a package. This is not implicit in the type
1 characters as such: type 1 interorbital
septa occur with all three types of palatines
and crania in the Elaeniinae and Fluvi-
colinae, and type 1 palatines occur with all
types of interorbital septa and cranial
types. The only correlation that appears
structural is that type 1 crania almost al-
ways occur with type 1 palatines; the
reverse, however, is not true. The uni-
formity in the Tyrannine cranium suggests
a late origin for this subfamily; the fact that
they are a highly successful and dominant
group with evidence of recent radiation
supports this view. On the other hand,
three of the character states delineating
this uniform cranium are primitive; only
the type 1 cranium is a derived state. This
leaves us with the anomaly of the most suc-
cessful subfamily being the most primitive
anatomically. Possibly the explanation is
that retention of the primitive or general-
ized characters left the Tyranninae with a
greater potential for radiation, which they
have only recently exploited.

SUMMARY

A new classification of the Tyrannidae is
presented, based on available morphologi-
cal, anatomical, behavioral and distribu-
tional data. The most useful recent studies
on the higher classification of the Tyran-
nidae were those of Warter ( 1965 ) on the
cranial characters of the Tyrannoidea, and
Ames ( 1971 ) on the syrinx of the Passeri-
formes. The family includes all those
genera accepted by Hellmayr (1927), five
genera formerly included in the Cotingidae
— Attila, Pseudattila, Casiornis, Laniocera
and Rhytipterna — and Corythopis from the
former family Conopophagidae.

The seven subfamilies of Hellmayr,
Fluvicolinae, Tyranninae, Myiarchinae,
Platyrinchinae, Euscarthminae, Serpo-
phaginae and Elaeniinae, are reduced to
three. The Fluvicolinae now include that
part of the Myiarchinae including the
genera Nuttallornis through OnychorJiyn-

Classification of Tyrant Flycatchers • Trcvl

y^>

chiis, and the remainder of the Myiarcliinae
plus the former Cotingid genera are
placed in the Tyranninae. The other four
hubfamilies are merged into one, for which
Elaeniinae is the oldest name. The sub-
famiUes are hsted in the order Elaeniinae,
Fkivicolinae, Tyranninae, since this seems
to represent the progression from most
primitive or generalized to most specialized
or successful.

The uenera are hsted within each sub-
family also from generalized to specialized.
However, these are such subjective deci-
sions that the order of Hellmayr (in
reverse) is not disturbed unless there is
positive evidence for making a change.
There has been Httle analysis of the genera
of flycatchers over the past century, and for
tlie most part they have been accepted un-
critically from checklist to checklist. Thirty-
six of Hellmayr's genera are here synony-
mized, one is resurrected and one new
genus is described. The latter is Zimmeriiis,
created for five species formerly in the
genus Tyranniscus. The new classification
is summarized in Appendix A.

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36. The Genera Elaenia and Myiopagis.
Amer. Mus. Novit., No. 1108, 23 pp.

. 1941b. Studies of Peruvian Birds, No.

37. The Genera Siiblegatus; Phaeomyias,
CciDiptostonia, Xantliomyias, PliyUomyias, and
Tyiaunisciis. Amer. Mus. No\it., No. 1109,
25 pp.

— . 1941c. Studies of Peruvian Birds, No.

38. The Genera Oieotriccus, TyiannuJus,
Acrochordopus, Oinithion, Leptopogon, Mio-
nectes, Pipromorpha and PyrocephaJiis. Amer.
Mus. Novit., No. 1126, 25 pp.

. 1955. Further notes on Tyrant Fly-
catchers ( Tyrannidae ) . Amer. Mus. Novit.,
No. 1749, 24 pp.

Sequence of Genera with Synonyms
Elaeniinae

PliyUomyias (syn: Xantliomyias, Oreotriccits,

Acrochordopus, and Tyranniscus nigrocapU-

lus, iiropygialis and cinereiceps)
Zimnicriiis (gen. nov. — inchides "Tyranniscus"

boUvianus, vilissijniis, cinereicapitla, graclli-

pes, viridiflavus)
Ornitliion (syn: Microtriccus)
Cainptostoma
Phaeomyias
Sid)Icgatus
Siiiriri
Tyrannuhis
Myiopagis
Elaenia
Mecocerctdtis
Serpopliaga
Inczia
Stigmatura

Anairetes (syn: Uromyias, Yanacea)
TacJiuris
Cidicivora
PoJystictus
Pseudocolopteryx
Euscarthmus

Mioncctcs (syn: Pipromorpha)
Leptopogon
PhyUoscartes (syn: Pogonotriccus, Leptotriccus,

Cajysicnipis)
Pseudotriccus (syn: Cacnotriccus)
Corythopis

Myiornis (syn: Perissotriccus)
Lopliotriccus (syn: Coloptertjx)
Atalotriccus

Poecdotriccus (syn: Taeniotriccus)
Oncostoma
Hcmitriccus (syn: Idioptilon, Euscaiihmornis,

SnctJdagaea, Microcochlearius, Cerato-

triccus)
Todirostrum
Cnipodectes
Ramplu)trigon
RJu/ncliocychis
Tohnomyias
Platyrinchus

Flu\ icolinae

OnycliorJiynchus

Myiotriccus

Terenotriccus

Myiobius

Myioplu)I)us

Aphanotriccus ( syn : Praedo )

Xenotriccus (syn: Aeclunolojylius)

PyrrlioDiyias

Mitrephanes

178 Bulletin Mu-scuin of Comparative Zoology, Vol. 148, No. 4

Contopus (syn: Nuttalluniis, Blaciciis)

Empidonax

Nesotricciis

Cncniotiicciis

Satjomis

PynxcpJiiiJus

Ochihocca (syn: Ochtlioniis, Culoiluimplius)

Mijiotheretes (syn: Cnemarchus, Ochthodiaeta)

Xohnis (syn: PyrofX')

Neoxolmis

Agriornis

MuscisaxicoJo

Lesson ia

Knipolegus (syn: Phaeotriceus, Entotriccus,

Entntiiohius)
Ihjincnops

Fluvicola (syn: Arundinicola)
Colonia

Alectrums (syn: Yeta))a)
Gubenietes
Satrapa
Tumhezia
Muscigndla
Hinindiiiea
Machetuinis
Miiscipipra

Tyranninae

Attila (syn: Pseudattila)

Casiornis

Rhytipterna

haniocera

Syristes

MyiarrJitis (syn: Ilylondx, Erihates)

Delturhyuclius

Pitangus

McgaihyiicJius

Myiozetctes

Conopias (syn: Coiyphotriccus)

Myiodynastes

Legatus

Eiyipidonomus

Tyniiniopsis

Tyraiunis (s>n: Miiscicora, Toliiuticlnts)

-St'dis Incertae
Xcnopsaris

APPENDIX B

Generic Changes Since Hellmayr (1927)

Acmchoidopus = PJiyJloniyias, here synonyniized
Acchniolophits Zinuiier 1938, Auk 55: 663 =

XcHutricctis, Wel)ster (1968: 289)
Ah'ctnirus - includes Yetapa, Short (1975: 269)
Auaiictcs 1850, replaces S})izitoniis 1920; Peters

and Griswokl (1943: 316); includes Ycnuicea

and Uromyids, which see
Aiilianotrirnis - includes Pnicdo, (Irisconi (1929:

176)

Arundinicola = Fhwicola, here synonymized
Attila - transferred from the Cotingidae, Snow

(1973: 7); Ames (1971: 155)
BlacicHS = Contopus, Bond (1943: 117)
Caenotriccus = Pseudotriccus, Zinuner (1940a:

22)
Capsicnipis = Phylloscartes, here synonymized
Casiornis - transferred from the Cotingidae, Snow

(1973: 7); Ames (1971: 155)
Ccnitotriccus = Idioptilon, Fitzpatrick (1976);

here synonymized with Hemitriccus
Cnemarchus = Myiotheretes, here synonymized;
see also Smith and Vuilleumier (1971: 193)
Colopteryx = Loplwtricctis, here synonymized
Colorhani])hus = Ochthoeca, here synonymized
Conopias - inchides Coryphotriccus, Meise ( 1949:

76)
Contopus 1855, replaces Myiochanes 1859, Ameri-
can Ornithologists' Union, Checklist Commit-
tee (1947: 449); includes Nuttallornis and
Blacicus, which see
Corythopis - added to the Tyrannidae, Ames,

Heimerdinger and Warter ( 1968 )
Coryjihotriccus = Conopias, Meise (1949: 76)
Elacnia - no longer includes Myiopagis, Zimmer

(1941a: 20)
Entotriccus — Knipolegus, Short (1975: 270)
Erihates - Myiarchus, Swarth (1931: 84)
Eumyiohius Brodkorb 1937, Proc. Biol. Soc. Wash-
ington, 50: 1 = Knipolegus, Zimmer, 1937b:
27
Euscartlonornis =z Idioptilon, Zimmer (1940a: 13)

=: Hemitriccus, here synonymized
Fluvicola - includes Arundinicola, here united
Hahrura 1859 = Polystictus 1850, Meyer de

Schauensee (1966: 374)
Hemitriccus - includes Ceratotriccus, Micrococh-
Icaritis, Sncthlagaea, Euscaiihniornis and
Idioptilon, here united; see also Fitzpatrick
( 1976 )
Hylonax = Myiarchus, Lanyon (1967b: 339)
Hymenops, 1828, replaces Lichenops, 1835, Meyer

de Schauensee (1966: 342) y

Idioptilon - includes Euscarthmornis, Zimmer
(1940a: 13) = Hemitriccus, here synony-
mized
Knipolegus - includes Phaeotriceus and Ento-
triccus, Short (1975: 270)
haniocera - transferred from the Cotingidae, Snow

(1973: 7); Ames (1971: 155)
Leptotriccus ^= Phylloscartes, here synonymized
Lichenops 1835 = Hymenops 1828, Meyer de

Schauensee (1966:" 342)
Lophotriccus - includes Colopteryx, here united
Micrococldearius — Idioptdon, Fitzpatrick (1976)

= Hemitriccus, here synonymized
Microtriccus = Ornithion, Zimmer (1941c: 3)
Mionectes - includes Pipromorpha, Dickey and

van Rossem (1938: 397)
Muscivora = Tyrannus, Smith (1966: 167)

Classification of Tyrant Flycatchers • Trcijlor

■7 -;.

Myiarchtis - includes Hijlonax and Erihatcs, which

see

Uyiochanes 1859 = Contopus 1855, American
Ornithologists' Union, Checklist Committee
(1947: 449)

Myiopagis - synonymized in Elaenia by Hellmayr,
but now recognized, Zimmer ( 1941a: 20)

Myiornis - includes Pcrissotiicciis, Zinnner {1940a:
220)

Myiotheretes - includes Cnemarchus and Ochtho-
diaeta, here imited; see also Smith and Vuil-
leumier (1971: 193)

Nuttalloinis = Contopus, Phillips, \hirshall and
Nhmson (1964: 90)

Ochthodiaeta = Myiotheretes, here synonymized;
see also Smith and X'uilleumier (1971: 193)

Ochthoeca - includes CoIoiJianiplius and Oehthor-
nis, here luiited

Ochthornis =: OeJithoeca, here synonymized

Oreotriectis = Pltyllomyias, here synonymized

Oinithion - includes Microtricciis, Zimmer ( 1941c:
3)

Peiissotiiccus = Myiornis, Zimmer (1940a: 22)

Pliaeotrieciis = Knipole^us, Short (1975: 270)

Pltyllomyias - includes Xanthomyias, Oreotriccus,
Aerochordopns and Tyranniscus (pt. ), here
united

Phylloscartes - includes Pogonotriccus, Lepto-
triccus and Capsiempis, here united

Pipromorpha z= Mionectes, Dickey and van Ros-
sem (1938: 397)

Pogonotriccus = Phylloscartes, here synonymized

Polystictus 1850, replaces Hahrura 1859 - Meyer
de Schauensee (1966: 374)

Praedo = Aphanotriccus, Griscom ( 1929 : 176 )

Pseudattila Zinnner 1936, Amer. Mus. Novit. No.
893: 6 = Attila, Short (1975: 265); trans-
ferred from the Cotingidae, Meyer de Schau-
ensee (1970: 297), Snow (1973: 7)

Pseudotriccus - includes Caenotriccus, Zimmer
(1940a: 22)

Pyrope = Xohnis; synonymized by Hellmayr
(1927: 10), resurrected by Meyer de Schauen-
see (1966: 335), and synon\aiiized again by
Smith and Vuillermiier (1971: 193)

Rhytiptcrna - transferred from the Cotingidae,
Snow (1973: 8); Ames (1971: 155)

Snethlagaea = Idioptilon, Fitzpatrick (1976) =
Hemitriccus, here synonymized

Spizitornis 1920 = Anairetes 1850, Peters and
Griswold (1943: 316)

Tolmarchus = Tyrannus, Meise (1949: 73); Bond
( 1958 )

Tyranniscus (nigrocapillus, uropygialis, cinereiceps)
= Phyllonnjias, here synonymized; the remain-
ing species are in the new genus Zinimerius.

Tyrannus - includes Muscivora and Tolmarchus,

which see
Uroinyias = Anairetes, Smith (1971: 275)
Xanthomyias — Phyllomyias, here synonymized
Xenopsaris - here placed at end of Tyraimidae,

incertae sedis
Xcnotriccus Dwight and Griscom 1927, Amer.

Mus. Novit. No. 254: 1 - includes Aechmolo-

phus, Webster (1968: 289)
Xolmis - includes Pyrope, Smith and Vuilleumier

(1971: 193)
Yanacea - Garriker, 1933, Proc. Acad. Nat. Sci.

Philadelphia 8S: 27 = Anairetes, here synon-
ymized
Yefapa = Alectrurus, Short (1975: 269)
Zimnterius - here described, includes Tyranniscus

holivianus, vilissimus, cinereicapilla, gracilipes,

and viridiflavus of Hellmayr

APPENDIX C

Ames' (1971: 158-162) Groups of Genera

1. Fluvicola group. The nucleus of this group is
composed of Xolmis, Neoxolmis, Agriornis,
Muscisaxicola, Fluvicola, Gubernetes, Knipo-
legus, Muscipipra and Phaeotriccus. Hymeno})s
is probably an offshoot. Satrapa and Ento-
triccus probably also belong here. Lessonia
agrees with them in musculatme.

2. Tyrannus group. The group includes Tyrannus,
Muscivora, Tohnarchtts, Tyrannoj)sis, Empido-
nomus, Myiodynastes, Megadnjnchus, Conopias
and Pitaugus (sulpliuratus only).

3. Nuttallornis group. Closest to the Fluvicola
group. Includes Nuttallornis, Sayornis, Cojito-
pus, Blacicus, Em})idonax, Aeehmolophits,
Aphanotriccus and possibly Muscigralla.

4. Myiohius group. Includes Myi()I)ius, Tereno-
triccus, Pyrrhomyias and Onychodiynchus. The
manakin Piprites probably also belongs here.

5. Myiarehus group. Includes Myiarchus, Eri-
hates, Attila, Casiornis, Lanioeera and Rhytip-
tcrna.

6. Colopteryx group. Contains Colopteryx, Onco-
stoma, Euscarthmornis, Hemitricctis, Myiornis
and Lophotriccus. Platyrinchus might belong
near here. Tolmonnjias, Rhynehocychis and
Platyrinchus differ greath- among themseKes.

7. Elaenia group. Included are Elaenia (including
Myiopagis), Suiriri, Camptostoma, Tyrannulus
and Phacomyias. Microtricciis and Tyranniscus
nigrocapillus probabK' belong near here.

APPENDIX D

Figures 2 and 3, and Table II of W'akter.
1965: 27-34

180 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

Type:

1 ^

'^>

Figure 2. Types of Nasal Septa. Left column, cross section; center column, ventral aspect; right column,
lateral aspect (all illustrations diagrammatic).

Classification of Tyrant Flycatchers • Traylor 181

Figure 3. Features of Type I Skulls. A, types of interorbital septa; B, types of palatines; C, types of crania
(posterior aspect).

182 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

Table 2. Varl\ble characters of flycatcher
SKtTLLS. Numbers in columns refer to "types"
illustrated in Figures 2 and 3. Column I —
Nasal septum. Column II — Interorbital sep-
tum. Column III — Palatines. Column IV —
Cranium. Column V — Palatomaxillaries ( +
= present).

Genus

III

FLUVICOLINAE

Agriomis 6

XoJmis 6

Mtiscisaxicola 6

Lcssonia 6

Mijiotheretes 6

NcoxnJmis 6

Ochthocca 6

Saijornis 6

Colonia 6

Guhentetes 6

Yciapa 6

Knipolegus 6

Entotricciis 6

Lichcuops 6

Fhtvicola 6

Anmdinicola 6

Fyroccphalus 6

Ochthornis 6

Muscigmlla 5

Satrapa 6

Machetornis 6

TYRANNINAE

Muscivora 1

Tymnntis 1

Empidonomus 1

Legatus 2

Sinjstcs 2

Myiodyuastes 1

Mcganjnchus 1

Coryphotriccus 1

Myiozctctes 1

Tyiannopsis 1

Pitangus 1

Tohnarchiis 1

MYIARCHINAE

Myiarchus

RJiytipterna

Eribatcs

Nesotriccus

Ntittallomis

Contopiis

Blacictis

Empidonax

2
2
2
2
6
6
6
6

2
2

1(4)

1(4)
2

2
3
3
3

3
2

1
1
1

2
1
1

3
2

2(3?)

2
3
3
3

3
1
2
2
3

+
+
+

+
+
+
+

+
+
+

+
+

Table 2. (continued)

Genus

III

Aechmolophtis 6

Cncnwtriccus 6

Mitrcphancs 6

Tcreiiotricciis 6?

Myiohius 6

Pynhoiuyias 6

MyiopJiobiis 6

Omjchorhynchus 3

PLATYRINCHINAE

Platyrinchus 3

Cnipodectes 1(3)

Tohuomyias 1(3)

Rliynchocychis 1(3)

EUSCARTHMINAE

Todirostrtnn

Oncostoma

Euscarthnioinis

SnctJdagaca

Lopliotriccus

Coloptcryx

Myiornis

Pogonotricciis

Leptotiicciis

Phylloscaites

Capshnnpis

Pscudocoloptcnjx

Hah rum

2
1

1?
1

1
3?

2?
2?
6

2
2

4?(1)

4
4
4
4

4
4
4
4
4
4
4
4
4
4
4
4
3

3(2)
2

o
O

3
3
3
2

2
2
2
3
2
2
2
1

3 3

1(2) 2

1 2

2(3) 2

2
1

1(2)

2(1)

SERPOPHAGINAE

?
5

Tachuris

Spizitornis

Stigmattira

Scr})op]}aga

Inczia

Mecocerculus

ELAENIINAE

2
5
5
5

4 2

4(3) 2(1)

4 1(2)

3
3
3
3
3
3
3
3
3
3
?

3
3

3
3
3
3

+
+
+

Myiopagis

3(2)

Elaenia

Suiriri

Sublegatus

Phaeomyias

Camptostoma

Xaniliomyias

3(2)

Tyrdiuiisciis

Tyraiiiiuhis

Microtriccus

2-3

Lcpto])ogon

Mionectes

Pipromorpha

Classification of Tyrant Flycatchers • Traijlor 183

INDEX

A.crochordopus 145
Aechmolophiis 161
/\griornis 143, 162
Alectrurus 164
Anairetes 150, 155, 173
Aphanotriccus 161
Arundinicola 143, 164
Atalotriccus 156
Attila 141ff, 166, 174

Blacicus 161

Caenotriccus 155
Camptostoma 148, 149, 173
Capsienipis 153
Casiornis 14 Iff, 166
Ceratotriccus 156, 157
Cnemarchus 162, 163
Cnemotriccus 161, 174
Cnipodectes 158
Colonia 164
Colopteiyx 156, 173
Colorhamphus 151, 163
Conopias 167, 169
Contopus 143, 161
Coryphotriccus 167
Corxthopis 1 56
Culicivora 151

Deltarh>nchus 141ff, 166

.Elaenia 148, 149, 155, 173
lEnipidonax 143, 161, 174
Eiiipidonomus 167, 169
Entotriccus 141, 143, 164
Erator 135
Eril)ates 141ff, 166
Euscarthniornis 156
Eusearthnius 151

Fluvicola 143, 162, 164

Gubernetes 143, 164

Hemitriccus 155, 156, 157
Hirundinea 164, 165
Hylonax 14 Iff, 166
Hymenops 164

ildioptilon 155, 156, 157
Inezia 150, 173

Knipolegus 143, 164

Laniocera 141ff, 166
Laphvctes 170
Legatus 167, 169, 174

Leptopogon 151, 155
Leptotriccus 153
Lessonia 164
Lichenops 164
Lipaugus 134
Lophotriccus 156, 159

Machetomis 164, 165
Mecocerculus 150, 153, 155, 173
Megarhynchus 167, 169
Microcochlearius 156, 157
Microtriccus 147
Mionectes 151
Mitrephanes 161, 174
Miiscigralla 162, 163, 164, 165, 173
Muscipipra 143, 164, 166
Muscisaxicola 143, 162
Muscivoia 167, 170
Myiarchus 14 Iff, 166, 174
Myiobius 143, 159, 160
Myiodynastes 167, 169
Myiopagis 148, 149, 150, 173
Myiophobus 143, 160, 174
Myiornis 156
Myiotheretes 162, 163
Myiotriccus 143, 160
Myiozetetes 167, 169

Neopelma 136
Neoxolmis 163
Nesotricciis 162
Nuttallornis 161, 174

Ochthodiaeta 163

Ochthoeca 141, 143, 163

Ochthornis 141, 163

Oncostoma 156, 157

Onychorhynchus 141, 143, 144, 153, 159, 173

Oieotricciis 145

Ornitbion 147

Oxvruncvis 133

135

156
148, 173

143, 164
145, 150, 155

153, 155

Pachyraniphus

Perissotriccus

Phaeomyias

Pliaeotricciis

Phyllomyias

Phxlloscartes

Pipiites 136

Pipromorpha 151

Pitangus 167, 169

Platvpsaris 135

Platyrinchus 144, 153, 158,

Poecilotiiccus 156, 157

Pogonotriccus 153

Polystictus 146, 151. 173

Praedo 161

174

184 Bulletin Museum of Comparative Zoology, Vol. 148, No. 4

Pseudatilla 141, 166

Pseudocolopteryx 151, 155, 173

Pseiidotricciis 155

P\TOcephalus 161

Pyrope 1 63

Pyrrhnnn ias 143, 160, 161, 174

Ramphotrigon 158

Rln nchocyclus 144, 158

Rhytipterna 141ft', 166

Satrapa 1 64
Sayomis 143, 161
Serpophaga 150, 170, 173
Snethlagea 156, 157
Stigniatura 150
Sublegatus 148, 149, 173
Suiriri 148, 149, 173
Sxristes 166

Tachuris 151

Taeniotriccus 143, 156, 157

Terenotricciis 160
Tityra 135
Todirostruni 144, 155,
Tolniarchus 167, 170
Tolmomyias 144, 158
Tunibezia 164
Tyranniscus 145
Tyrannopsis 167, 170
Tyrannulus 148, 173
Tyranniis 167, 170

Uromyias 150

Xanthomyias 145
Xenopsaris 135
Xenotriccus 161
Xolmis 143, 162, 163

Yanacea 150
Yetapa 143, 164

Zimmerius 147, 173

156, 157

us ISSN 0027-4100

bulletin OF TH

seum

Comparative
Zoology

The Orb-Weaver Genera Metepeira,

Kaira and Aculepeira in America

North of Mexico (Araneae: Araneidae)

HERBERT W. LEVI

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 5
29 NOVEMBER 1977

PUBLICATIONS ISSUED

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chusetts, 02138, U.S.A.

'it

THE ORB-WEAVER GENERA METEPEIRA, KAIRA AND
ACULEPEIRA IN AMERICA NORTH OF MEXICO
(ARANEAE: ARANEIDAE)

HERBERT W. LEVI'

Abstract. Metepeiia and Kaira are known only abundant in semidesert, an unusual habitat

in the Americas. Eleven species of Metepeira live
nortli of Mexico, frequently in semiarid regions.
A number of species are sympatric in the south-
western United States and are often collected to-
gether. Only four species of Kaira have been found
north of Mexico; in one of these only females are
known, in another, only males. The habits of Kaira
species remain a mystery. Aculepeira is mainly
an Eurasian genus, but two species live in North
America; both North American species also occur
in Siberia. One, A. packardi, is found in the
Arctic, in western mountains, in meadows and in
sagebrush. Tlie other, A. carhonarioides, has been
discovered only between boulders in talus slides
of the Arctic, the Rocky Mountains, the Gaspe
Peninsula and the White Mountains of New
Hampshire. Identification of Eurasian species was
difficult because few specimens were available.

INTRODUCTION

for orb-weavers. The two Aculepeira spe-
cies are found in mountain meadows and
tahis slopes where it is very hot during the
day, but becomes freezing at night, even
in midsummer. But here the similarity ends.
After careful study I found Metepeira and
Kaira are not closely related to Araneus
and are Hmited to the Americas, but Acule-
peira is an Old World genus close to
Araneus. Two Siberian species of Acule-
peira are believed to have spread over the
North American continent into various "va-
cant" niches. At least four other species
are known from Eurasia (Figs. 187-231),
but no others are known from America.

For this study the collections of the Mu-
seum of Comparative Zoology (MCZ) were
The genera sequence of these revisions used. I would like to thank the following
is unfortunately not phylogenetic; instead for specimens: J. A. Beatty; D. E. BLxler;
their arrangement develops from the order J. E. Carico; R. Crabill (National Museum
in which specimens could be sorted out of Natural Histoiy); C. D. Dondale (Ca-

most easily from the available collections.
I started with Argiope and tlie large species
of Araneus. This paper includes species
that have at times been confused with those
in Araneus and other genera. When I started
this revision of the three genera, I thought
Aculepeira was close to Metepeira. Species
of both genera have a median wliite streak
on the venter of the abdomen, the median
apophysis of the male palpus has two fla-
gella and species of both inliabit extieme
climates. The Metepeira species are most

^ Museum of Comparative Zoology.

Bull. Mus. Comp. ZooL, 148(5

nadian National Collections, CNC); S. I.
Frommer (University of California, River-
side Collections); M. Grasshoff (Sencken-
berg Museum, Frankfurt, SMF); J. Gruber
( Naturhistorisches Museum, Wien); N.
Horner; M. Hubert (Museum National
d'Histoire Naturelle, Paris ) ; W. R. Icenogle;
B. J. Kaston; J. B. Kethley (Field Museum);
T. Kronestedt (Natural History Museum,
Stockholm, NRS); R. Leech; W. B. Peck
(Exline-Peck Collection); W. J. Gertsch
and N. I. Platnick (American Museum of
Natural Historv, AMNH and Cornell Uni-

185-238, November, 1977 185

1<S6 Bulletin Mu-sciini of Comparative Zoology, Vol. 148, No. 5

versity Collections, CUC); G. Levy (He-
brew University Collections, JerLisalem,
HUCJ); S. Riechert; V. D. Roth; R. X.
Schick and P. H. Arnaud (California Acad-
emy of Science); W. A. Shear; W. T. Sedg-
wick; K. Thaler (KT); M. E. Thompson;

E. S. Sutter ( Naturhistorisches Museum,
Basel; NMB); B. Vogel; H. K. Wallace;

F. R. Wanless (British Museum, Natural
History, BMNH); W. Star^ga (Pohsh Acad-
emy of Sciences); and H. V. Weems (Flor-
ida State Collection of Arthropods). In
addition, V. D. Roth and W. R. Icenogle
supplied living specimens from Arizona and
California, and W. R. Icenogle provided
careful habitat notes on sympatric Califor-
nia Metepeira species. P. Brignoli (PBC),
besides loaning specimens, made a gift of
a specimen and provided helpful informa-
tion. M. Troeger mapped the species and
reworded awkward sentences, D. Randolph
typed the various stages of the manuscript.
The observations on the American Acule-
peira species were made while at the Rocky
Mountain Biological Laboratory, Crested
Butte, Colorado; supported in part by Na-
tional Science Foundation Grant No. G-2335
in 1956 and 1958. The research and publi-
cation was supported in part by National
Science Foundation grant number DEB
76-115568.

METHODS

The method used to obtain the results
presented here is the usual one: comparing
specimens or those moiphological features
of specimens that have proven most useful
in the past in taxonomy of spiders (Levi,
1977). The useful morphological charac-
ters, in all spiders, are the genitalia (the
female epigynum, the male palpus); in
araneids, the color patterns of the carapace
and abdomen and the proportions and seta-
tion of the legs. Differences of several char-
acters between groups of specimens are
important because they indicate popula-
tions that do not interbreed. Often numer-
ous outline drawings were made to find
such differential characters.

Species differences proved subtle in
Metepeira; these species are difficult to
separate. Not only are several species sym-
patric, but no differences in their habits
are even known at present. Unlike Mete-
peira species, the two American species of
Aculepeira are quite variable, with no two
specimens exactly alike and some specimens
more than twice the size of others. Despite
this variability, no populations could be
segregated as having several distinct fea-
tures. There is the possibility, though, that
the eastern population of A. carbonarioides
is distinct. A larger series of specimens is
needed to ascertain the differences. But
even if they are not distinct moiphologi-
cally, can we be certain that the four isolated
populations of A. carbonarioides are po-
tentially interbreeding? In an attempt to
answer this, I tried to raise Colorado speci-
mens, but could not keep them alive in the
laboratory.

Some of the problems of taxonomy are
illustrated by the Eurasian species. Can I
be sure that the three Siberian species de-
scribed in the past ( but with no specimens
now available except for a female from
Kamchatka) are the same species as those
I did examine that were found in the Amer-
ican Arctic? While the study of specimens
of the three Mediterranean species (Figs.
187-217) indicated they were far apart mor-
phologically and easy to separate, the few
specimens close to A. armida available
from Asia were puzzling. Some that were
labeled A. karabagi (Karol) seemed very
distinct. But further specimens from Israel
(one from Galilee and others from the
Negev Desert) were intermediate with A.
armida, having the characteristic lamellae
posterior of the epigynum bent at right
angles (Fig. 211) but much wider than the
illustrated specimens from Europe. Despite
this I believe A. karahagi to be a distinct
species with the oldest name A. noseki.
Drensky (1943) synonymized A. victoria
Thorell with A. armida. Can I be sure by
examining two specimens labeled Araneus
victoria in the available collections that

Metepeira, Kaira, Aculepeira ' Levi 187

Drensky was correct? I suspect Drensky
was wrong, but it is impossible to obtain
large series of these populations. Perhaps
A. ormido is a highly variable species and
all doubtful specimens belong to it. I de-
cided then to leave the Old World prob-
lems to a colleague closer to the critical
areas (the Balkans and Asia Minor) who
can resolve the question by further collect-
ing and field observations.

I Metepeira F. P. -Cambridge

Metepeira F.P. -Cambridge, 1903, Biologia Cen-
trali-Americana, Araneidea, 2: 457. Type spe-
cies by original designation M. spinipes F.P.-
Cambridge, 1903. The name is feminine.

Note. In 1942 Chamberlin and Ivie
named 14 species of Metepeira and men-
tioned 16 others in a paper on diverse new
species from the Americas. It is difficult
to see this as a revisionary study since the
majority of the University of Utah collec-
tions then available had not been deter-
mined (except for some immatures!). Ap-
parently, individual specimens that looked
different were picked out of the collection
and named. No distinguishing features
were given by Chamberlin and Ivie for
the new species and only a few of the illus-
trations feature diagnostic characters. The
usual wording in the description regarding
diagnosis is "showing minor differences in
the palpus," or "the palpus and epigynum
are distinct as shown by the figures." Nev-
ertheless, most of the names of Chamberlin
and Ivie actually do apply to different spe-
cies and only a few names are synonymized.

Diagnosis. Metepeira differs from other
araneid genera in having the eye region
lighter than the remainder of the carapace
(when in alcohol). The posterior head re-
gion is often darkest (Figs. 3, 37, 39).
Unlike most other araneid genera, Mete-
peira has a median, longitudinal white line
contrasting with the black background on
the venter of the aljdomen (Plates 1, 2, 3;
Figs. 4, 38, 40). This line is absent, how-
ever, in the light-colored M. gosoga, in
which only two black streaks remain (Fig.

36 ) . There are usually white spots on black
background on each side of the spinnerets
(Plates 1, 2; Figs. 4, 38, 40). Unlike all
other genera, the white line continues on
the sternum (Plates 1, 2; Figs. 4, 38, 40).
The few exceptions to this are M. foxi, M.
grandiosa and M. datona (Figs. 86, 95, 98,
100). In still another difference between
Metepeira and most other araneid genera
related to Araneus, the combined length of
metatarsus and tarsus is longer than that
of the patella and tibia of the same leg
(Fig. 3). (An unusual exception is M.
datoiui; Plate 4.) An additional diagnostic
trait is the very small, weakly sclerotized
epigynum (Fig. 14) and the small palpus
that has a median apophysis bearing two
filiform appendages (flagella) (Figs. 10,
19, 20, 26, 27). The palpal tibia has two
strong macrosetae (Figs. 8, 10, 19), as does
the palpal patella (Fig. 8) in most species.
The web, too, is diagnostic (see below),
differing distinctly from that of species in
related genera.

Metepeira, like Aculepeira, has a ventral
median white mark on the abdomen, and
the median apophysis of the male palpus
has two flagella. Metepeira differs from
Aculepeira by having a more spherical ab-
domen (Figs. 3, 4) and by the relatively
smaller genitalia with a different structure.
Metepeira is close to Kaira but the Mete-
peira abdomen is spherical (Figs. 3, 4);
the Kaira abdomen is higher than long with
tuljerculate dorsal humps (Figs. 122-126)
and the ventral abdominal mark is indis-
tinct.

Description. The carapace has the an-
terior of the head lighter and, in all species,
it is covered by white down ( Plates 1, 2, 3).
The thoracic depression is an indistinct lon-
gitudinal shallow groove. In all species the
anterior median eyes are slightly larger or
subequal in size with the others ( Figs. 1,5).
Also in all species, the anterior median eyes
are separated from each other by their
diameter or slightly more, and from the lat-
erals by one to one and one-half diameters
(Figs. 1, 5). The posterior median eyes are

188 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Plate 1. Metepeira labyrinthea. Upper left, female (Florida). Righit, retreat, in barrier-web, and orb (Indiana);
photo B. Open, Bottom, orb with barrier-web and retreat, cornstarch dusted (Tavernier, Florida). Horizontal
diameter of viscid area of web, 23 cm.

Metepeira, Kaira, AcuLEPEiRA • Levi 189

slightly less than their diameter apart in
females, and two to three diameters from
the laterals ( Fig. 1 ) . In males the pos-
terior median eyes are their radius to
slightly less than their diameter apart, and
one and one-half to slightly more than two
diameters from the laterals (Fig. 5). The
height of the clypeus is less than the diam-
eter of the anterior median eyes. The en-
dites are longer than in many other araneid
genera ( Fig. 2 ) . The legs are banded
(Plates 1, 2; Fig. 3). The abdomen is
usually oval, but spherical in M. foxi and
M. grandiosa (Figs. 94-100) and is wider
than long in M. datona ( Plate 3; Fig. (S5 ) .
The abdomen always has a dorsal folium
pattern, which is more distinct posteriorly,
and there may be black shoulder patches
anterior-Iaterally (Plates 1, 2; Figs. 3, 68,
85). The black venter, with its character-
istic white streak (Plate 1), contrasts with
the light dorsum.

Living Metepeira labyrinthea, M. grin-
nclli and M. crossipes, as well as M. foxi
and M. datona that I have seen and photo-
graphed, have on the abdomen, especially
to the sides of the folium, some red pigment
that washes out in alcohol. Metepeira grin-
nelli has reddish areas on the borders of
the white line on each side of the folium,
and anteriorly the folium has black pig-
ment grading into brown. There is also a
reddish brown area on the venter on each
side of the black mark.

Males have no hook on the first coxa and
no groove on the second femur. The male's
first femur has strong macrosetae anteriorly,
with the largest medially in the M. foxi
group (Fig. 96), and distally in other
groups (Figs. 12, 13). The fourth coxae
are not modified. While males are smaller
than females, in most species they do come
in various sizes — some small, some larger,
the larger ones presumably having gone
through more instars and a longer period
of growth.

The proportions of most species are about
the same and the sizes overlap, but mem-
bers of the M. foxi group are, in general.

Plate 2. Metepeira grinnelll, female (Arizona).

slightly smaller. Measurements of a female
M. labyrinthea from Virginia are: total
length 7.0 mm; carapace 2.8 mm long, 2.2
mm wide. First femur, 3.8 mm; patella
and tibia, 3.9 mm; metatarsus, 3.6 mm;
tarsus, 1.2 mm. Second patella and tibia,
3.1 mm; third, 1.9 mm; fourth, 2.9 mm.
Measurements of a male from Virginia are:
total length 4.2 mm; carapace 1.9 mm long,
1.6 mm wide. First femur, 3.3 mm; patella
and tibia, 3.2 mm; metatarsus, 3.4 mm;
tarsus, 1.1 mm. Second patella and tibia,
2.6 mm; third, 1.3 mm; fourth, 1.8 mm.

190 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Plate 3. Metepeira crassipes (Riverside Co., California)
female venter; right, male. Laboratory photographs.

Upper photographs, females, dorsal. Lov\/er left,

Genitalia. The epigyiuim is very small
and lightly sclerotized, making it difficult
to study (Fig. 14). The openings are in a
species-characteristic depression (Fig. 18).
Because of the small size of the soft struc-
ture, it is necessary, when the species is
tmcertain, to examine it as a temporarily
cleared microscope mount. Below the open-
ing is a sclerotized spherical structure from
which ducts lead to the seminal receptacles
(Figs. 18, 25, 32).

The femur of the male palpus lacks the
pointed tooth usually present on the prox-
imal end in all related genera and also lacks
the tooth of the endite that is usually facing
that of the femur. The median apophysis
of the palpus bears two flagella ( Figs. 10,
20) and often has a keel distally (ventrally
on the palpus) (Figs. 10, 26, 27, 91, 93,
109, 111). Even though the median apo-
physis is the most prominent part of the
palpus, its shape varies and, therefore, it

Metepeira, Kaira, Aculepeira • Levi 191

is only occasionally of diagnostic value. The
shape of the triangular conductor (Figs.
10, 11) is similar in many species, but is
wider than long in M. foxi (Fig. 92). Of
greatest importance for diagnosis is the
embolus, which is hidden partly behind
the flagella of the median apophysis and
partly by the overhanging terminal apo-
physis (Figs. 9, 10, 19, 26). The portion
of the embolus containing the duct has a
soft lobe hanging above it in some species
(Figs. 9, 10, 20, 27, 34). Because of its
diagnostic importance, it is unfortunate that
this lobe is partly hidden. In virgin males
the embolus is capped (Figs. 34, 52). The
cap has been lost in presumably mated
males (Figs. 33, 51). The cap, which has
a minute barb at its tip, is found in the
opening of the epigynum, plugging it (Figs.
25, 73 ) , and apparently preventing a second
mating of the female. There is never more
than one cap found on each side in the
epigynum. This cap contains an open duct
in Metepeira. Some palpi (when handled
through different solutions) emit tube-
shaped material from their tips, probably
propelled outward by osmotic pressures.
Below the section of the embolus carrying
the duct is a larger lobe in all species that
projects on the outside of the conductor
(right in the left palpus) in ventral view
(Figs. 9-11) in the contracted palpus.
Large and small males of the same species
may have slightly different proportions in
the softer parts of the palpus, but not in
the embolus.

Natural History. Metepeira has an orb
and a barrier-web — an irregular web to the
side and slightly above. Lubin (1975) re-
cently suggested that the barrier web may
be a moisture gathering device. The orb
may be incomplete on top (Plate 1). The
hub has a mesh and several trap lines travel
from the hub to the retreat in the center of
the barrier- web (Plates 1, 5). The retreat
is a small cap of silk covered by insect re-
mains or leaves (Plates 1, 5). The web of
M. labyrinthea is often in dead branches.
Egg-sacs are brown, lenticular and hung in

Plate 4. Metepeira datona, female (Florida).

a string below the retreat with the most
recent on the bottom (Comstock, 1940;
Kaston, 1948 ) . According to Kaston ( 1948 ) ,
M. labyrinthea has about 35 radii and there
are 5 to 6 egg-sacs with up to 63 eggs. The
cocoon lasts until spring. Metepeira in
Riverside County, California observed by
Icenogle (personal correspondence) feed,
to a great extent, on crane flies. Comstock
claims that M. labyrinthea matures in fall,
but in the collections there are mature males
and females gathered from early summer
to fall. Judging by Metepeiras abundance
in semidesert areas, the dense retreat and
barrier-web must provide sufficient pro-
tection against desiccation. In many habi-
tats it is the only orb-weaver found. One
Mexican and Central American species, M.
spinipes F.P.-Cambridge, is colonial, the
others are not.

The Metepeira species of the eastern
states are known to have very different
habitat preferences: M. labyrinthea usually
places its webs in shrubs of deciduous
forests; M. grandiosa pahistris selects north-

192 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Plate 5. Orb-web, barrier-webs and retreats of Metepeira grandlosa alpina from Colorado. Upper left, side
view, sprayed with Krylon'R; white paint. Upper right, side view, dusted with cornstarch. Bottom, frontal view
from slightly below, dusted with cornstarch. Diameter of viscid area of lower web about 25-30 cm.

Metepeira, Kaira, Aculepeira • Levi 193

ern bogs; and M. datona uses shaded shnibs
on ocean shores. One might tlins expect
the numerous western species to each have
different habitats. But four species (M.
crassipes, M. ventura, M. foxi and M. g.
grandiosa) have all been found side by side
in California buckwheat {Eriogonum fas-
cicidattim) and sage (Artemisia calif or-
nica) in Riverside County, California. The
first two species are commonly found in this
environment, according to the best docu-
mented collections by W. Icenogle.

All species are commonly preyed upon
by mud-dauber wasps of the genera Try-
poxijlon and Trypargiltim (Sphecidae).

Species. No Metepeira species are known
from other continents, all are American,
and they range from Alaska to Tierra del
Fuego. All, at one time or another, have
been referred to as M. labyrinthea because
of their similar size and coloration. ( Mete-
peira foxi and M. datona are smaller than
most species.) Roewer (1942) gives the
distribution of M. labyrinthea from Pata-
gonia to Labrador; Bonnet (1957) claims
it extends from Canada to Patagonia, Chile,
as well as appearing in east Africa. Actually
it has a very limited distiibution, occurring
only in the eastern United States ( Map 1 ) .
Systematic characters. Numerous char-
acters were checked and illustrated in the
hope of finding discontinuities or of finding
characters that would go together, thus in-
dicating species. I studied absolute sizes,
proportions, coloration and color pattern,
and the ventral and anterior macrosetae of
the male femora. Color pattern is useful to
segregate the M. gosoga and the M. foxi
group of species. More macrosetae are pres-
ent in large specimens of each species than
in small ones, so I did not find these setae
very useful (Figs. 12, 13); however, males
of the M. foxi group (M. datona, M. foxi,
M. grandiosa) have the longest macrosetae
in the middle of the femur (Fig. 96), while
the M. labyrinthea has them distally (Fig.
I 6). Various aspects of the epigynum, dif-
I ferent ones in different species, are useful.
The shape of the terminal apophysis of the

palpus segregates the M. lalnjririthea and
M. foxi species groups (Figs. 19, 91). The
conductor, because it is too similar and
variable, is not of much use. The median
apophysis, which is sometimes very distinct,
is variable and therefore of doubtful use
alone, but when used with the shape of the
embolus it proves a satisfactory character.
Matching males with females of the same
species can be a problem. It is helpful to
collect males with females because this fa-
cilitates identifications, in some species the
male, in others the female, is easiest to
determine.

Species groups. The species north of
Mexico belong to two species groups: M.
labyrinthea and M. foxi. The M. laby-
rinthea group has a longitudinal white line
across the sternum (Fig. 4); the median
apophysis of the male palpus has only a
short distal keel beyond the flagella-bearing
proximal part (Figs. 10, 19, 20, 26, 27).
All species within this group are about the
same size. Species in the M. foxi group
have a black sternum (Fig. 95) and the
median apophysis of the palpus has a distal
(ventral on the palpus) tuberculate keel
(Figs. 91, 93, 104, 105, 115, 116). The spe-
cies in this group are smaller in size. Two
of the three species of the M. foxi group
are common and come in large numbers
in collections. It is interesting that of the
species of the M. labyrinthea group, whose
sternal white line is often broken and partly
missing (Fig. 38), M. grinnelli also has an
extension (though relatively small) on the
median apophysis beyond the flagella
(Figs. 26, 27). It is difficult at present to
decide which of these species groups is
the derived and which the more primitive.
I consider M. datona most primitive be-
cause of the shorter metatarsus and the less
specialized coloration.

Metepeira grandiosa is puzzling and I
consider all specimens to be M. grandiosa
with three subspecies for the three distinct
allopatric forms. I have a suspicion, how-
ever, that with more data, some will turn
out to be distinct.

194 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

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Metepeira, Kaira, Aculepeira • Levi 195

Key to female Metepeira north of Mexico

1. Sternum black (Figs. 95, 98, 100),

sometimes in Florida specimens with
a lighter brown anterior and posterior

patch (Fig. 86)

Sternum with a white longitudinal
band (Figs. 4, 36, 40), rarely broken

and partly missing (Fig. 38) 6

2(1) Abdomen wider than long, anterior
half of dorsum light (Plate 4; Fig. 85 h
openings of epigynum on each side in
depression (Fig. 78); Florida coast

datona

Abdomen spherical to slightly longer
than wide, usually with a dorsal folium
(Figs. 94-100); opening of epigynum
hidden (Figs. 87, 101-112); most of
area but not in southeastern United

States 3

3(2) Coxae black like sternum (Fig. 98);
Canada, south to Maine and South

Dakota (Map 2) grandiosa paltistris

Coxae light yellowish to orange 4

4(3) Epigynum witli a bordered, longitu-
dinal depression on each side of narrow
scape ( Fig. 87 ) ; western United States

(Map 2) foxi

- Epigynum with transverse depression

(Figs. 101, 106, 112); North Dakota,

Colorado, western United States 5

5(4) Epigynum scape wide at base (Fig.
106); in posterior view middle piece
shorter \'entrally than lateral ones ( Fig.
107); western Canada to Oklahoma;
Chihuahua west to eastern Oregon

(Map 2) grandiosa alpina

Epigynum scape usually narrow, some-
times with a slight median ridge ( Fig.
112); in posterior view middle piece
about same lengtli ventrally as lateral
ones (Fig. 113); British Columbia to
California (Map 2) _- grandiosa grandiosa
6(1) Epigynal scape fleshy triangular, great-
est width as wide or wider than visible
base on either side (Figs. 41-43); de-
pressions on each side of scape small,
visible diameter less than diameter of
their rim (Fig. 41); western Texas,
Chihuahua, to California and Gulf of

California (Map 1) - arizonica

Scape otherwise an equal to or narrower
tlian base visible to side of it ( Figs. 14,
21, 47); depressions small or larger

(Figs. 47, 53, 61) 7

7(6) Visible depression on each side of
scape appearing as anterior-posterior
slits with their lateral borders parallel
(Fig. 70); southern Texas to central
Mexico, West Indies (Map 1) -- minima

Depression otherwise (Figs. 14, 28,
47) 8

8(7) Posterior rim of depression narrow
(Fig. 61) and a round opening visible
on each side in posteroventral view
( Fig. 62 ) ; Texas, New Mexico to Chi-
huahua (Map 1) - - comanche

- Epigynum otherwise (Figs. 14, 53) .— 9

9(8) Eastern North America to western
Oklahoma, western Texas (Map 1);
epigynum as in Figs. 14-18 „ lahijrinthea
Western United States, New Mexico

to Pacific coast 10

10(9) Carapace light brown, with posterior
head dark ( Fig. 35 ) and whitish abdo-
men with two ventral black streaks
(Fig. 36); epigynum as in Fig. 28;
Utah, Nevada, Arizona to California

(Map 1) gosoga

Carapace, except for eye area, dark
brown (Figs. 39, 59) and venter of
abdomen black with median white lon-
gitudinal line ( Figs. 40, 60 ) ; epigynum
otherwise H

11(10) Width of scape less tlian one-fourth
width of epigynal base ( Fig. 53 ) ; pos-
terior rim of depression narrow ( Fig.
53); California, Sonora and Baja Cali-
fornia (Map 1) Ventura

Width of scape about one-third width
of epigynal base (Figs. 21, 47); pos-
terior rim of depression wider (Figs.
21, 47); Arizona, Oregon to Cali-
fornia 12

12(11) Depressions round and small with
width of posterior border wider tlian
visible diameter of depression (Fig.
47); Cahfornia to Baja California

(Map 1) crassipes

- Depression flaring and width of its
posterior border narrower tlian visible
diameter of depression (Fig. 21); Ari-
zona to Oregon and California, Chihua-
hua and Sonora (Map 1) grinnelli

Key to male Metepeira north of Mexico

1. Sternum black, without longitudinal
light line (Figs. 86, 95, 98, 100); base
of palpal embolus at tlie distal tip of
bulb (Figs. 82, 91, 109); median apo-
physis with a prominent ventrally di-
rected tubercular keel beyond base of
two flagella (Figs. 84, 93, 111) 2

Sternum with a median longitudinal
light band (Figs. 4, 36, 38); terminal
apophysis o\erhanging palpal embolus
(Figs. 10, 11, 19, 26); median apophy-
sis witliout such a keel or with only ji
small smooth extension (Figs. 21, 27,
34, 46) 6

196 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

2(1) Eml)()liis tliick as in Figs. 82, 84; ter-
minal apophysis a narrow, curved, soft
pronu (Fig. 83); Florida coast datona
Enil)()lus thin (Figs. 91, 104, 105, 109,
111, 115, 116); terminal apophysis
otherwise (Figs. 92, 104, 109, 110,
115); whole region except southeastern
United States 3

3(2) Coxae black like sternum (Fig. 98):
Canada, south to Maine and South
Dakota (Map 2) grandiosa palustris

— Co.xae light \ellowish to orange; west-
ern North America 4

4(3) Embolus of palpus with thin terminal
part pointed at 45° angle to wider basal
portion (Figs. 91, 93); tubercular keel
of median apophysis larger than prox-
imal flagella bearing part (Figs. 91,
93 ) ; western United States ( Map 2 ) ._ foxi

— Embolus of palpus with terminal part
cur\'ed at right angle to wider basal
portion (Figs. 104, 105, 109, 111, 115,
116); tubercular keel of median apo-
physis e(iual to or smaller than prox-
imal flagella bearing part (Figs. 105,
111, 116); North Dakota, Colorado and
western United States 5

5(4) Terminal part of embolus narrower
(Fig. Ill); western Canada to Okla-
homa, Chihuahua, west to eastern Ore-
gon, eastern California (Map 2)

lJ.mndio.sa alpina

— Terminal part of embolus wider ( Fig.
116); British Columbia to California

( Map 2 ) grandiosa grandiosa

6( 1) Embolus of palpus with a parallel lobe
above terminal portion containing duct
(Figs. 20, 27, 34); lobe partly hidden
by terminal apophysis (Figs. 19, 26,
33) 7

- Embolus of palpus without such a lobe
(Figs. 46, 52, 58, 67) 9

7(6) Embolus tip strongly curved, the lower
edge of its base concave ( Fig. 20 ) and
median apophysis with an indistinct
small keel beyond flagella ( Figs. 19,
20); eastern United States to western
Oklahoma and western Texas ( Nhip 1)
labyrintJica

- Embolus tip less curved (Figs. 27, 34)
and median apophysis with larger keel
(Fig. 27) or no keel; western United
States 8

8(7) Embolus curved and lobe overhanging
tip (Fig. 27); median apophysis with a
keel beyond flagella (Figs. 26, 27);
Arizona to Oregon and California, Chi-
huahua and Sonora ( Map 1 ) giinnclli

- Embolus tip with upper edge straight

and lobe less than half length of tip
(Fig. 34); median apophysis without
keel (Fig. 34); Utah, Nevada, Arizona

to California ( Map 1 ) gosoga

9(6) Proximal f lagellum of median apophysis
( left one of left palpus ) more than four
times length of distal one ( Figs. 74,
75); southern Texas to central Mexico,
West Indies (Map 1) minima

— Proximal flagellum of median apophysis
less than tliree times length of distal
one (Figs. 46, 58); Texas, California
and southwestern United States 10

10(9) Base of embolus with a distinct lobe at
base of tip ( Figs. 66, 67 ) ; Texas, New
Mexico to Chihuahua ( Map 1 ) _ conmnche

- Embolus otlierwise or if with lobe; not
from Texas or New Mexico 11

11(10) Embolus gracefully cuwed, fairly long
( Figs. 45, 46 ) ; base of median apophy-
sis wide and both flagella recurved
(Figs. 45, 46); western Texas, Chihua-
hua to California and Gulf of California
(Map 1) arizonica

- Embolus curved but shorter (Figs. 52,
58 ) ; base of median apophysis narrow,
distal flagellum not recurved (Figs.
52, 58); Oregon to Baja California,
Sonora 12

12(11) Proximal flagellum of median apophy-
sis, noticeably longer and much wider
than transparent distal one (Figs. 57,
58); median apophysis higher (Figs.
57, 58 ) ; California, Sonora and Baja
California ( Map 1 ) - Ventura

— Proximal flagellum of median apophysis
almost subequal in length and width to
distal one (Figs. 51, 52); median apo-
physis narrower (Figs. 51, 52); Cali-
fornia to Baja California ( Map 1 )

___-—_, crassipes

Metepeira tabyrinthea (Hentz)

Plate 1; Figures 1-11, 14-20; Map 1

Epeira Jabyrinthea Hentz, 1847, J. Boston Soc.
Natur. Hist, 5: 471, pi. 31, fig. 3, $. Type
specimens from North Carolina and Alabama,
destroyed. Emerton, 1884, Trans. Connecticut
Acad. Sci., 6: 314, pi. 34, fig. 8, pi. 36, fig. 11,
9, S. Keyserling, 1893, Spinnen Amerikas, 4:
215, pi. 10, fig. 160, 9, £. Emerton, 1902,
Common Spiders, p. 174, figs. 408-410, $, web.

Epeira crucifera Keyserling, 1864, Sitzungsber.
Naturwiss. Gesell. Isis, Dresden, p. 132, pi. 6,
figs. 11, 12, $. Female holotype from Baltimore
in till' British Museum, Natural History, exam-
ined. (The locality of Baltimore is not pub-

Metepeira, Kaira, Aculepeira • Levi 197

Map 2. Distribution of Metepeira datona, M. grandiosa and M. loxi.

lished Iwt is on the label in the vial. Name
preoccupied by Lucas, 1835. )
Metepeira lahyrinthea: — F. P. -Cambridge, 1903,
Biologia Centrali-Americana, Araneidae, 2: 458,

pi. 43, figs. 6, 7, $, <^. Comstock, 1940, The
Spider Book, rev. ed., p. 476, figs. 187, 476-479,
? , c5 , web. Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. ser., 7(1): 63, figs. 161-164,

198 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

2, $. Kaston, 1948, Bull. Connecticut Geol.
Natur. Hist. Surv., 70: 226, figs. 704, 724, 2036,
$, $, web.
Aranca keyserlingi Roewer, 1942, Katalog dei
Araneae, 2: 861. New name for Epeira cnicifcm
thought preoccupied. NEW SYNONYMY.

Variation. Females are from 4.0 to 8.6
mm total length, carapace 1.7 to 3.5 mm
long, 1.1 to 3.0 mm wide. Males are 3.0 to
6.8 mm total length, carapace from 1.6 to
3.0 mm long, 1.2 to 2.2 mm wide. The
length of the first patella and tibia is 1.2 to
1.3 times carapace length in females, 1.6
times in males.

Didii^uosis. Over most of its range, M.
labijrinthea is the only Metepeira. In the
north its range overlaps with M. grandiosa
palustris and in Florida with M. datona;
both, unlike M. lahijrinthea (Fig. 4), lack
the longitudinal light line on the sternum.
Only in Texas does M. lahijrinthea overlap
the range of other species, and females can
be separated by the scape of the epigynum,
which has a narrow neck at its base. The
scape is wider (Figs. 14, 15) than that of
M. minima and M. comanche and narrower
than that of M. arizonica, and the epigynal
depression on each side of the scape (Fig.
18) is a different shape from those of the
three other species. The embolus, unlike
these three other species, is strongly curved
with a lobe overhanging it (Figs. 19, 20).

Natural History. This is a forest species
and is found on shrubs. It has been col-
lected in floodplain forest in Tennessee, in
thick forest on trees and bushes in Virginia,
on the edge of oak wockIs and fields in Wis-
consin, in oak, hickory and birch woods in
Missouri, in a deciduous forest in Kansas, in
woods in Texas and in the shade below
trees growing among cacti in the Florida
Keys. Adult males are found during July
and August, females from spring to late fall.

and, in southern Florida, females are found
all year. '

Distribution. Massachusetts, southern On-
tario, southern Wisconsin, to western Okla-
homa and western Texas, south to the Flor-
ida Keys and Tamauhpas (Map 1).

Metepeira grinnelli (Coolidge)

Plate 2; Figures 21-27, 37, 38; Map 1

Epeira labijrinthea grinnelli Coolidge, 1910, J.
Entomol. Zool., Claremont, 2: 281. There are
no type specimens in the Los Angeles County
Museum, Pomona College or in the major arach-
nid collections.

Metepeira douglasi Chamberlin and Ivie, 1941,
Bull. Univ. Utah, biol. ser., 6(3): 18, figs. 21-
23, $ . Female holotype from Santa Ana, Cali-
fornia in the American Museum of Natural His-
tory, examined. Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. sen, 7(1): 66, figs. 169-170,
c5. NEW SYNONYMY.

Note. Females of this species are often
larger than those of M. lahijrinthea, the col-
oration is more pronounced and the banding
of the legs is more distinct as described by
Coolidge. Also it is the most common spe-
cies of the three localities cited by Coolidge:
Palo Alto, Pasadena and Lompoc.

Variation. Most specimens have the ven-
tral white line of the sternum broken by
black pigment (Fig. 38); sometimes the an-
terior or posterior part of the line is missing.
Total length of females 5.0 to 9.4 mm, cara-
pace 2.3 to 4.1 mm long, 1.7 to 3.4 mm wide.
Total length of males 3.6 to 6.7 mm, cara-
pace 1.9 to 3.3 mm long, 1.4 to 2.6 mm wide.
The first patella and tibia is 1.2 to 1.4 times
the carapace length in the female, 1.5 to 1.7
in the male.

Diagnosis. This species tends to be larger
in size and darker in coloration than sym-
patric species. It is very close to M. lahij-
rinthea. Females differ by the longer scape,

Figures 1-11. IVIetepeira labyrinttiea (Hentz). 1-4. Female; 1. Eye region and chelicerae. 2. Lateral. 3.
Dorsal. 4. Ventral. 5-11. Male: 5. Eye region and chelicerae. 6. Left femora, ventral. 7. Dorsal. 8. Left
cymbium, tibia and patella; bulb removed. 9. Left median apoptiysis, conductor and embolus, pulled apart.
10. Left palpus, expanded. 11. Left palpus, ventral view.

Figures 12, 13. M. arizonica Chamberlin and Ivie, left male femora, ventral view: 12. (Chiricahua tVlountains,
Arizona.) 13. (Canyon Lake, Maricopa County, Arizona.)

Metepeira, Kaira, Aculepeira • Levi 199

«§^«\

Abbreviations, a, terminal apophysis; c. conductor; e, embolus; h, hematodocha; m, median apophysis; y,
cymbium.

Scale lines. Figs. 1-7, 12-13, 1.0 mm; Figs. 8-11, 0.1 mm.

200 Bulletin Musew7i of Comparative Zoology, Vol. 148, No. 5

which does not have the neck of that of
M. la])yrintheo. Also, in a posteroventral
view of the epigynum, the openings appear
in two circnlar dark areas (Fig. 22), those
of M. Ial)i/rint]ica in a dark streak (Fig. 15).
M. grinnelli differs from M. arizonica in
liaving the posterior head region darker
nsnally tlian tlie thorax to the sides (Fig.
37). While most females can readily be
separated from M. arizonica by the much
narrower scape and rims (Fig. 21), and
from M. Ventura by the wider scape and
wider rims, they can be distinguished as
well by the pockets of the median depres-
sion (Fig. 25) that are visible ventrally
using a cleared posterior view. The epigy-
num differs from that of M. crassipes by
the longer scape and larger depression on
each side (Fig. 21). The embolus of the
male is not as strongly curved as that of M.
Jalyijrinthea (Figs. 26, 27) and its over-
hanging lobe is larger (Fig. 27). Most dis-
tinct is the ventral extension, or keel, of the
median apophysis that extends beyond the
two flagella ( on the right of the left palpus,
Figs. 26, 27) and is less distinct in M.
hihyrintliea. This keel is variable in shape.

Natural History. Adult males are found
from July to September, females from July
to October; in Sonora females are found in
April. None of the specimens in collections
come with ecological data.

Distribution. Arizona, Oregon, Califor-
nia to Chihuahua and Sonora ( Map 1 ) .

Metepeira gosoga Chamberlin and Ivie
Figures 28-36; Map 1

Metepeira gosoga Chamberlin and Ivie, 1935, Bull.
Univ. Utah, biol. ser., 2(8): 21, figs 82-83, $.
Female holotype from Pilot Knob Valley, Mo-
have Desert [34 km west of Johannesburg, San
Bernardino County], California in the American
Museum of Natural History, examined.

Variation. Total length of females 7.5 to
9.4 mm, carapace 3.2 to 4.1 mm long, 2.4
to 3.2 mm wide. Total length of males 4.4
to 5.8 mm, carapace 2.3 to 2.9 mm long, 1.7
to 2.4 mm wide. The first patella and tibia
of females is 1.1 to 1.2 times the carapace
length, that of the male 1.4 times.

Diagnosis. Metepeira gosoga can readily
be recognized by the light colored carapace
having only the posterior head region dark,
by the white abdomen, which only pos-
teriorly shows a folium (Fig. 35), and by
the two adjacent ventral black marks sur-
rounded by white pigment (Fig. 36). The
middle piece of the epigynum in posterior
view is sclerotized at its dorsal end ( toward
the abdomen ) but not ventrally toward the
short stubby scape. In cleared posterior
view the ventrolateral pockets each have a
median extension ( Fig. 32 ) that is not pres-
ent in the related species of M. grinnelli, M.
arizonica and M. laljyrinthea. The embolus
of the male has a lobe above ( Fig. 34 ) , but
unhke that of M. grinnelli and M. lahy-
rinthea, the lobe is shorter than the sclero-
tized part bearing the duct (Fig. 34).

Natural History. Adult males and fe-
males have been collected from June to Au-
gust. The only specimen with notes was
collected on a cholla cactus ( Opuntia sp. ) .
It is likely that this species is often found on
desert vegetation.

Distribution. Southern Utah to southern
Arizona west to California (Map 1). One
specimen from Mount Tamalpais State
Park, near Bootjack, Marin County, Califor-
nia, 28 Nov. 1968 (M. Bentzien) was con-
sidered doubtful and is not mapped.

Metepeira arizonica Chamberlin and Ivie
Figures 12, 13, 39-46; iVIap 1

Metepeira arizonica Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 69, figs. 182-
187, 9 , $ . Female holotype, paratypes from
Canyon Lake, W 111° 30' : N 33° 30', Arizona
[Tonto National Forest, 25 mi. east of Phoenix,
Maricopa Co.], in the American Museum of Nat-
ural History, examined.

Variation. Rare specimens are light in
color on the carapace and dorsum. Total
length of females 4.7 to 8.4 mm, carapace

2.0 to 3.5 mm long, 1.5 to 2.3 mm wide.
Total length of males 2.7 to 3.8 mm, cara-
pace 1.5 to 1.9 mm long, 1.2 to 1.6 mm wide.
The first patella and tibia of the female is

1.1 to 1.2 times the carapace length, that of
the male 1.3 to 1.4.

Metefeira, Kaira, Aculepeira • Levi 201

Figures 14-20. Metepeira labyrinthea (Hentz). 14-18. Epigynum: 14. Ventral. 15. Posteroventral. 16. Pos-
terior. 17. Lateral. 18. Posterior, cleared. 19. Left male palpus, mesal. 20. Median apophysis and embolus.

Figures 21-27. M. grinnelli (Coolidge). 21-25. Epigynum. 21. Ventral. 22. Posteroventral 23. Posterior. 24.
Lateral. 25. Posterior, cleared. 26. Male palpus, mesal. 27. Median apophysis and embolus.

Scale lines. 0.1 mm.

202 Bulletin Museum of Comparative Zoology, Wol. 148, No. 5

Diagnosis. Unlike M. grinnelli and M.
gosoga, but like M. crassipes, the carapace
is very dark brownish black except for the
contrasting light transverse anterior head re-
gion (Fig. 39). The epigynum of M. ori-
zonica has a swollen-looking appearance,
and unlike all related species, the diameter
of the scape is wider than the width of the
base on each side of the scape (Figs. 41-44).
Metepeira crassipes is very similar, though
slightly smaller. The epigynum has a simi-
lar small depression with seemingly swollen
lips; the scape, however, is much narrower
and has a proximal constriction. The grace-
fully curved embolus of the male (convex
below and lacking a lobe above (Figs. 45,
46) is longer than that of M. ventura. The
two flagclla of the median apophysis ( Figs.
45, 46) are about the same width, unlike
those of M. ventura, and are directed more
posteriorly than those of M. crassipes. In ad-
dition, the median apophysis ( Fig. 46) lacks
the ventral keel beyond the flagella that is
present in M. grinnelU. The sternal light
band is never broken (Fig. 40), while in
M. ventura it often is broken.

Natural History. Adult males have been
collected from March to November, females
from March to December. Several speci-
mens from Yuma County, Arizona came
from alfalfa fields, one came from a cactus
in Organ Pipe National Monument; others
were from oak-pine- juniper woodland in
Cave Creek Canyon, Chihuahua Movmtains,
Arizona.

Distribution. Western Texas, northern
Arizona to California to Chihuahua and
Baja California (Map 1).

Metepeira crassipes Chamberlin and Ivie
Plate 3; Figures 47-52; Map 1

Metepeira josepha Chamberlin and Ivic, 1942, Bull.
Univ. Utah, biol. sen, 7(1): 64, fig. 165, ?.
Female holotype from Kings Mtn. near Palo Alto,
California in the American Museum of Natural
History, examined. NEW SYNONYMY.

Metepeira crassipes Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 66, figs. 171-
173, $ , S • Male holotype, female, male para-

types from Laguna Beach, California in the
American Museum of Natural History, examined.

Note. Although M. josepha has a page
priority, as first revisor I prefer to use M.
crassipes since the male is the type and
several specimens are available from the
typo locality.

Variation. The white sternal band may
be broken as in M. grinneUi. The height of
the base of the median apophysis is variable.
Total length of females 4.9 to 7.2 mm, cara-
pace 2.2 to 3.2 mm long, 1.7 to 2.3 mm wide.
Total length of males 2.9 to 4.7 mm, cara-
pace 1.6 to 2.3 mm long, 1.2 to 1.7 mm wide.
The first patella and tibia is 1.0 to 1.2 times
the carapace length in females, 1.1 to 1.3
times in males.

Diagnosis. Although sizes overlap, this
species is noticeably smaller than M. grin-
neUi when collected with it. Females can
be separated from M. grinneUi by the shorter
scape having a slight constriction (Figs.
47, 49) and from both M. grinneUi and
M. ventura by the smaller depression on
each side of the scape (Fig. 47). The
females differ from M. arizonica in that
the side of the base of the epigynum, which
is visible on each side of the scape, is
wider than the scape itself (Fig. 47).
While females can be confused with M.
arizonica, males are more likely to be mis-
taken for M. ventura. Differences, however,
can be detected. Males of this species have
both flagella of the median apophysis sub-
equal in length (Figs. 51, 52), while those
of M. ventura are quite unequal. The space
surrounded by the proximal, posteriorly
directed flagellum and by the base of the
median apophysis is equal or larger than
the area of the median apophysis ( below it
in Figs. 51, 52), while it is smaller in M.
ventura. The embolus has a much shorter
distal duct-bearing portion (Figs. 51, 52)
than that of M. arizonica, and the terminal
apophysis (Fig. 51) appears to be smaller
than that of M. ventura.

Natural History. Males have been col-
lected from April to October. A large num-
ber of specimens were collected from Cali-

Metepeira, Kaira, Aculepeira • Levi 203

;...■£;-

'^"^^

/Too

Figures 28-36. Metepeira gosoga Chamberlin and Ivie. 28-32. Epigynum: 28. Ventral. 29. Posteroventral. 30.
Posterior. 31. Lateral. 32. Posterior, cleared. 33. Left male palpus, mesal. 34. Median apophysis and
embolus. 35. Female, dorsal. 36. Female, ventral.

Figures 37, 38. M. grinnelli (Coolidge). 37. Female, dorsal. 38. Female, ventral.

Figures 39, 40. M. arizonica Chamberlin and Ivie. 39. Female, dorsal. 40. Female, ventral.

Scale lines. 0.1 mm except Figs. 35-40, 1.0 mm.

204 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

fornia buckwheat (Eriogonnm fasciculatum)
and sage {Afiemisia cciUfornica) in River-
side County, California.

Distribution. California and Baja Califor-
nia (Map 1).

Metepeira ventura Chamberlin and Ivie
Figures 53-60; Map 1

Metepeira ensenada Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 65, figs. 166-
168, $ . Male holotype from beach near En-
senada [Baja California], Mexico in the Ameri-
can Museum of Natural History, examined.
NEW SYNONYMY.

Metepeira ventura Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 67, figs. 175-
179, ? . Female holotype, 1 male and 3 female
paratypes from between Oxnard and Santa
Monica, California, in the American Museum of
Natural History, examined.

Note. Although M. ensenada has page
priority, I prefer to use the name M. ventura
for this species, since numerous specimens
were available from the type locality of M.
ventura that are more characteristic of this
species.

Variation. A median light streak may be
present on the carapace (Fig. 59) and the
light sternal band is sometimes broken. Fe-
males are from 4.7 to 7.4 mm total length,
carapace 2.4 to 2.9 mm long, 1.6 to 2.4 mm
wide. Males are 2.6 to 6.0 mm total length,
carapace from 1.5 to 3.0 mm long, 1.0 to
2.3 mm wide. The length of the first patella
and tibia is 1.0 to 1.3 times carapace length
in females, 1.4 to 1.6 in males. There is
some variation in the width of the epigynal
scape.

Diagnosis. The middle of the posterior
head region often has a light streak not
found in M. grinnclli, M. crassipes or M.
arizonica, but this is not always present.
The light sternal band may be broken, un-
like that of M. arizonica. The epigynum
of M. ventura usually has a much narrower
scape (Figs. 5.3-56) than the sympatric M.
crassipes, M. grinnelli, M. arizonica and
M. gosoga, and the depression to the sides
of the scape are wider with the posterior
rim narrower (Fig. 53) than that of the

other species. The median depression of
the epigynum faces the scape but there are
pockets on each side as seen in a cleared
epigynum from the posterior view (Fig.
56). The embolus of the palpus lacks the
lobe above the part containing the duct
(Fig. 58), just like that of M. arizonica and
unHke that of M. grinnelli. The embolus is
less gracefully curved and shorter than that
of M. arizonica. The two branches of the
median apophysis are more slender than
those of M. arizonica. In addition, the
proximal one is large, the distal one small
(Figs. 57, 58), unlike those of M. crassipes.
The space surrounded by the proximal,
posteriorly directed flagellum and median
apophysis is less than the area of the median
apophysis (below it in Fig. 57), while it
is equal or larger in M. crassipes.

Natural History. Adult males have been
collected from May to July, females from
May to September. One collection of this
species from Lompoc, California came
from a mustard (Brassicaceae) field, an-
other from manzanita (ArctostapJiylos)
chaparral, and a larger collection from
California buckwheat (Eriogonum fa.scicu-
Jafum) and California sage { Artemisia cali-
fornica).

Distribution. California, Sonora, Baja
California (Map 1).

Metepeira comanche new species
Figures 61-69; Map 1

Type. Male holotype from 9.7 km west
of O'Brien, Haskell Co., Texas, 3.II.1971,
from the annual legume guar ( Cyamopsis
tetragonolopa) , C. E. Rogers, in the Mu-
seum of Comparative Zoology. The name is
a noun in apposition after the Indian tribe
of the Texas plains.

Description. Female from Webb Co.,
Texas. Anterior light area of head grades
gradually into dark area posteriorly (Fig.
68). A median light streak in many speci-
mens just anterior to thoracic depression. I
Legs banded. Dorsum of abdomen with '
usual folium; posteriorly, venter often with
white transverse bar connecting with longi- !

Metepeira, Kaira, Aculepeira • Levi 205

Figures 41-46. Metepeira arizonica Chamberlin and Ivie. 41-44. Epigynum: 41. Ventral. 42. Posterior. 43.
Lateral. 44. Posterior, cleared. 45. Left male palpus, mesal. 46. Median apophysis and embolus.

Figures 47-52. M. crassipes Chamberlin and Ivie. 47-50. Epigynum: 47. Ventral. 48. Posterior. 49. LateraL
50. Posterior, cleared. 51. Male palpus, mesal. 52. Median apophysis and embolus.

Scale lines. 0.1 mm.

206 Bulletin Muscinn of Comparative Zoolo^ij. Vol. 148, No. 5

tudinal mark (Fig. 69). There is one .spot
on each side, anterior to the spinnerets.
Posterior median eyes 0.(S diameter of an-
terior medians, anterior laterals 0.7, pos-
terior laterals 0.8 diameter of anterior
median eyes. Anterior median eyes slightly
less than their diameter apart, 1.3 from
laterals. Posterior median eyes their diam-
eter apart. Total length 6.5 mm. Carapace
2.6 mm long, 2.2 mm wide. First femur,
2.9 mm; patella and tibia, 3.2 mm; meta-
tarsus, 2.9 mm; tarsus, 0.5 mm. Second
patella and tibia, 2.7 mm; third, 1.6 mm;
fourth, 2.2 mm.

Male from Wells Co., Texas. Posterior
median eyes 0.7 diameter of anterior medi-
ans, anterior laterals 0.7 diameter, posterior
laterals 0.6 diameter of anterior medians.
Anterior median eyes 0.6 diameter apart,
slightly less than their diameter from later-
als. Posterior median eyes 0.5 diameter
apart, two diameters from laterals. Anterior
margin of chelicerae with four teeth, first
one and then three, spaced close together;
posterior margin with three teeth. Total
length 4.7 mm. Carapace 2.3 mm long, 1.7
mm wide. First femur, 3.2 mm; patella and
tibia, 3.4 mm; metatarsus, 3.2 mm; tarsus,
1.1 mm. Second patella and tibia, 2.8 mm;
third, 1.6 mm; fourth, 2.2 mm. The male ho-
lotype from Haskell Co., Texas: total length
3.0 mm. Carapace 1.6 mm long, 1.2 mm
wide. First femur, 2.5 mm; patella and
tibia, 2.3 mm; metatarsus, 2.1 mm; tarsus,
0.8 mm. Second patella and tibia, 1.9 mm;
third, 1.0 mm; fourth, 1.4 mm.

Variation. No two males are the same
size and even males collected together
seemed to vary tremendously in size. With
this variation in size come differences in the
macrosetae: the smaller males have fewer.
There are also differences in the palpus of
small and large specimens but not in the
shape of the sclerotized median apophysis
and embolus. The white sternal line may
be broken. Females are from 4.5 to 7.2
mm total length, carapace 2.2 to 3.2 mm
long, 1.8 to 2.4 mm wide. Males are 3.2
to 5.2 mm total length, carapace 1.8 to 2.4

mm long, 1.5 to 1.9 mm wide. The length
of the first patella and tibia is 1.1 times the
carapace length of females, 1.3 to 1.6 times
that of males.

Diapio.sis. Female M. comanche differ
from the sympatric M. la])yrinthea by hav-
ing a much wider depression on each side
of the scape with a narrower rim to the
sides and posteriorly (Fig. 61). The area
adjacent to the scape and part of its base
is wide, extending posteriorly in the de-
pression (Figs. 61, 62). The embolus of the
male (Figs. 66, 67) lacks the overhanging
lobe present in M. labyrintliea and differs
from that of M. arizonica by being more
evenly curved and by having a short upper
lobe, which is not overhanging (Figs. 66,
67). The color of the venter of the abdo-
men is often light, showing a transverse
light mark anterior to the spinnerets (Fig.
69).

Natural History. Adult males and fe-
males have been collected from May to No-
vember. None of the collections come with
habitat data.

Distribution. Texas, New Mexico, Chi-
huahua (Map 1).

Metepeira minima Gertsch
Figures 70-77; Map 1

Metepeira luinimd Gertsch, 1936, Amer. Mus. No-
vitates, no. 852: 10, fig. 31, S. Male holotype
from Edinburg, Texas in the American Museum
of Natural History, examined. Chamberlin and
Ivie, 1942, Bull. Univ. Utah, biol. sen, 7(1): 67,
fig. 174, ?.

Metepeira jamaieensis Archer, 1958, Amer. Mus.
Novitates, no. 1922: 16, fig. 33, $. Female
holotype from Port Henderson, St. Catherine
Parish, lamaica, in the American Museum of
Natural History, examined. NEW SYNONYMY.

Variation. Females are from 4.5 to 6.5
mm total length, carapace 1.9 to 2.6 mm
long, 1.4 to 2.2 mm wide. Males are 2.6 to
4.2 mm total length, carapace from 1.3 to
2.7 mm long, 0.9 to 1.6 mm wide. The
length of the first patella and tibia is 1.1
times carapace length in females, 1.4 times
in males.

Diag,nosis. The female M. minima epigy-

Metepeira, Kaira, Aculepeira • Levi 207

Figures 53-60. Metepeira ventura Chamberlin and Ivie. 53-56. Epigynum: 53. Ventral. 54. Posterior. 55. Lat-
eral. 56. Posterior, cleared. 57. Left male palpus, mesal. 58. Median apophysis and embolus. 59. Female,
dorsal. 60. Female, ventral.

Figures 61-69. M. comanche n. sp. 61-65. Epigynum; 61. Ventral. 62. Posteroventral. 63. Posterior. 64.
Lateral. 65. Posterior, cleared. 66. Male palpus, mesal view. 67. Median apophysis and embolus. 68. Fe-
male, dorsal. 69. Female, ventral.

Scale lines. 0,1 mm except Figs. 59, 60, 68. 69, 1.0 mm.

208 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

niim has a narrow scape with a parallel slit-
like depression on each side ( Fig. 70 ) that
is quite different from that of other North
American species. Males, too, can be
readily separated from other species be-
cause the proximal flagellum of the median
apophysis is several times as long as the
distal one ( Figs. 74, 75); in all other species
north of Mexico the two flagella are of
about equal length.

Natural Histonj. Adults can be found in
all seasons. No observations of this spe-
cies are available from the United States or
Mexico. In Jamaica I have observed adults
in webs along a road in a shady situation
and away from the shore.

Distribution. Southern Texas, to central
Mexico and Jamaica (Map 1).

Metepeira datona Chamberlin and Ivie
Plate 4; Figures 78-86; Map 2

Metepeira datona Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. sen, 7(1): 68, fig. 196, $.
Female holotype from Daytona Beach, Florida in
the American Museum of Natural History, ex-
amined.

Metepeira inerma Bryant, 1945, Bull. Mus. Comp.
Zool., 95: 378, fig. 20, 9 . Female holotype from
Cap Haitien, Haiti in poor physical condition in
the Museum of Comparative Zoology, examined.
NEW SYNONYMY.

Description. Female from Flagler Co.,
Florida. Eye region of carapace orange;
and orange longitudinal median line ( Fig.
85). Lateral eyes in dark bro\\'n area. Ster-
num black-brown, sometimes with lighter
brown area anteriorly and another posteri-
orly (Fig. 86). Legs are yellow- white with
nanow, dark broken bands on dorsum.
Dorsiun of abdomen light anteriorly with
a black posterior folium. Sides black dis-
tinctly delineated towards dorsum, but not
towards venter. Venter with a median
white .spot on black. Posterior median eyes
subequal to anterior medians. Anterior
and posterior laterals 0.8 diameter of
anterior median eves. Anterior median
eyes L2 diamc>lers apart, 1 diameter from
laterals. Posterior median eyes slightly less

than their diameter apart, 2.5 from laterals.
Abdomen is wider than long ( Plate 4; Fig.
85). Total length 3.9 mm. Carapace 1.8
mm long, 1.5 mm wide. First femur, 2.2
mm; patella and tibia, 2.3 mm; metatarsus,
1.5 mm; tarsus, 0.6 mm. Second patella and
tibia, 1.9 mm; third, 1.0 mm; fourth, 1.7
mm.

Male from the Bahama Islands. Color-
ation as in female. Abdomen, dorsum,
however, more conservatively colored with
a median dorsal line from anterior to pos-
terior. Anterior light patches surrounded
by black with a posterior dark patch ex-
tending more anteriorly (Plate 4; Fig. 85).
Posterior median eyes 0.8 diameter of an-
terior medians, laterals 0.7 diameter. An-
terior median eyes slightly more than their
diameter apart, the same distance from lat-
erals. Posterior median eyes 0.6 diameter
apart, 1.5 from laterals. No tooth on base
of palpal femur or on endite. Two femoral
macrosetae and two on tibia. Abdomen
longer than wide. Total length 3.2 mm.
Carapace 1.8 mm long, 1.5 mm wide. First
femur, 3.2 mm; patella and tibia, 3.5 mm;
metatarsus, 2.4 mm; tarsus, 0.9 mm. Second
patella and tibia, 2.7 mm; third, 1.3 mm;
fourth, 1.8 mm.

Variation. A male from southern Florida
had the eye region the same color as the
thoracic area. Females are from 2.6 to 4.6
mm total length, carapace 1.2 to 1.8 mm
long, 1.0 to 1.6 mm wide. Males are 2.0 to
3.2 mm total length, carapace 1.0 to 1.8 mm
long, 0.9 to 1.5 mm wide. The length of
the first patella and tibia is 1.1 to 1.3 times
the carapace length in females, 1.9 times
in males. The sternum is brownish black
and sometimes has a lighter area in the an-
terior and posterior sections.

Diagnosis. Unlike other Nortii American
species, M. datona has an abdomen that is
wider and more spherical than it is long;
the anterior of the dorsum has a light patch
framed by black (Figs. 85, 86). It differs,
of course, from small Araneus species of a
similar color, by the ventral median longi-
tudinal white stripe on the abdomen (Fig.

Metepeira, Kaira, Aculepeira ' Levi 209

Figures 70-77. Metepeira minima Gertsch. 70-73. Epigynum: 70. Ventral. 71. Posterior. 72. Lateral. 73.
Posterior, cleared. 74. Left male palpus, mesal. 75. Median apophysis and embolus. 76. Female, dorsal. 77.
Female, ventral.

Figures 78-86. M. datona Chamberlin and Ivie. 78 81. Epigynum: 78. Ventral. 79. Posterior. 80. Lateral.
81. Posterior, cleared. 82-84. Male palpus: 82. Mesal. 83. Ventral. 84. Median apophysis and embolus. 85.
Female, dorsal. 86. Female, ventral.

Scale lines. 0.1 mm except Figs. 76, 77, 85, 86, 1.0 mm.

210 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

(S6). The epigyniim is very different from
that of other species because the openings
appear as small slits on each side of the
wide depression (Fig. 7S). The palpus dif-
fers from that of M. foxi by having the
embolus heavier and wider (Fig. 82) and
by having the terminal apophysis a soft
projecting arm (Figs. 82, 83).

Nattiral Historij. In Florida adult females
have been collected only from November to
March. Metepeiia datona has been col-
lected from vegetation on the beach, from
shore shrubs and from beach grape ( Cocco-
loha uvifem) in Florida and Jamaica. An
adult male wa.s collected in mangroves on
Saddle Bunch Keys, in June.

Distribution. Florida coast and Greater
Antilles (Map 2).

Metepeira foxi Gertsch and Ivie
Figures 87-96; Map 2

Mcfcpciia foxi Gertscli and Ivie, 1936, Amer. Mus.
Novitates, no. 858, p. 20, figs. 42-44, 9, S-
Male holotype and female paratype from Rich-
field, Utah, in the American Museum of Natural
History, examined. Chaml)erlin and Ivie, 1942,
Bull. Univ. Utah, Iwol. ser., 7(1): 71, figs. 197,
198, 9, S ■ Roewer, 1942, Katalog der Araneae,
1: 869. Bonnet, 1957, Bibliographia Araneorum,
2: 2820.

Metepeira naneUa ChamlierHn and Ivie, 1942, Bull.
Univ. Utah, biol. ser., 7(1): 71, fig. 199, $.
Female holotype and paratype from Fillmore,
Utali, in the American Museum of Natural His-
tory, examined. NEW SYNONYMY.

Variation. Females are from 3.6 to 6.1
mm total length, carapace 1.7 to 2.2 mm
long, 1.2 to 1.6 mm wide. Males are from
2.7 to 4.5 mm total length, carapace 1.6 to
4.2 mm long, 1.3 to 1.7 mm wide. The
length of the first patella and tibia is 1.2 to
1.4 times carapace length in females, 1.5
to 1.7 times in males.

Dia<i,no.sis. Metepeira foxi differs from
all other species north of Mexico, except
M. '^randiosa and M. datona, in lacking a
white longitudinal line on the .sternum. It
does not occur in Florida, as does M. datona,
and females can be distinguished from M.
'^randiosa by the narrow scape of the epigy-
1 in II I, which has a bordered longitudinal de-

pression visible on each side (Fig. 87).
Male M. foxi differ from those of M. grandi-
osa by the 45" angle of the embolus (Figs.
91, 93); in M. grandiosa the bend of the
embolus is about 90". Metepeira foxi males,
in addition, have a conductor that is higher
(Fig. 92) than that of M. grandiose.

Natural History. Males have been col-
lected from March to July, most commonly
in April and June. Metepeira foxi has been
collected by sweeping at the edge of ponde-
rosa pine (Finns ponderosa) woods and in
meadows in Colorado, from meadows in
Wyoming, on sagebrush (Artemisia sp.) in
California and Oregon, from alkali marsh
vegetation in California and on saltbush
(Atriplex sp.) and hilaria (Hilaria sp.) in
Utah. Specimens are commonly collected
together with M. grandio.sa; their habitats
must be similar.

Distribution. Alberta to New Mexico
and west to British Columbia and Cali-
fornia (Map 2).

Metepeira grandiosa Chamberlin and Ivie
Plate 5; Figures 97-116; Map 2

Epeira lahyrintliea — "bog variety": — Emerton,
1915, Trans. Connecticut Acad. Sci., 20: 138,
fig. 6, 9, $.

Metepeira grandiosa Chamberlin and Ivie, 1941,
Bull. Univ. Utah, biol. ser., 6(3): 17, figs. 24-
26. Female holotype from Ben Lomond, Cali-
fornia, in the American Museum of Natural His-
tory, examined.

Metepeira palomara Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 72, figs. 200-
204, 9 , c? . Female holotype and paratypes from
Mt. Palomar, California, in the American Mu-
seum of Natural History, examined. NEW
SYNONYMY.

Metepeira dakota Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. ser., 7(1): 73, figs. 205-207,
9 , (5 . Male holotype and female paratype from
Noonan, North Dakota, in the American Mu-
seum of Natmal History, examined. NEW
SYNONYMY.

Metepeira palitstris Chamberlin and Ivie, 1942,
Bull. Univ. Utah, biol. ser., 7(1): 73, figs. 208-
210, 9, c^ . Female holotype from Divide
Count V, Nortli Dakota, in the American Museum
of Natural History, examined. NEW SYNON-
YMY.

Metepeira (il})ii\a Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. ser., 7(1): 74. Female holo-

Metepeira, Kaira, Aculepeira ' Levi 211

Figures 87-96. Metepeira foxi Gertsch and Ivie. 87-90. Epigynum: 87. Ventral. 88. Posterior. 89. Lateral.
90. Posterior, cleared. 91-93. Left male palpus: 91. Mesal. 92. Ventral. 93. Median apophysis and embolus.
94. Female, dorsal. 95. Female, ventral. 96. Left male femora, ventral.

Figures 97, 98. M. grandiosa palustris Chamberlin and Ivie, juvenile (Nova Scotia). 97. Dorsal. 98. Ventral.

Figures 99, 100. M. grandiosa alpina Chamberlin and ivie, female (northern Arizona). 99. Dorsal. 100. Ventral.

Scale lines. 0.1 mm except Figs. 94-100, 1.0 mm.

type and nunierou.s female paratypes from Fish is so distinct that 1 use .siib.specific names;

Lake, Utah, in American Museum of Natural palustris for the eastern and northern, al-

History, examined. NEW SYNONYMY. '^. r u c l at ^ ■ l . .w-^

■^' pina tor the Rocky Mountani and grandiosa

Subspecies. The variation of this species for the Pacific subspecies. Evidence for

212 Bulletin Museum of Cuiitparative Zoology, Vol. 148, No. 5

all belonging to one species is the follow-
ing. No consistent differences could be
found in the genitalia, although many draw-
ings of the epigyna, the internal female
genitalia, the embolus and the conductor of
the palpus were made. Most distinct is
M. grandiosa paliistris, which has black
coxae (Fig. 9(S). A collection from Medi-
cine Hat, Alberta includes a female with
black coxae, a male with yellow coxae; from
Stockade Lake, Custer County, South Da-
kota one female has black coxae, another
yellow; from Greeley County, Kansas a
female has yellow coxae with some dark;
from Medicine Hat, Alberta one female has
black coxae with orange distal rings; all
are border localities. Several collections
from East Rosebud Canyon, Carbon
County, Montana, 1963 to 1967 (Vogel col-
lection) are also intermediate: large fe-
males, small males, variously marked coxae,
some all black, some with two coxae black,
two yellow on each side and some with
coxae having yellow and black marks.

Diagnosis. Unlike most Metepeira, but
like M. foxi, the median longitudinal band
of the sternum is missing, the sternum is
entirely black (Figs. 98, loO). But unHke
M. foxi, the epigynum has a transverse de-
pression on each side posteriorly ( Figs. 101,
106, 112) in which the openings are located
laterally (Figs. 103, 108, 114 )^ The palpal
embolus is evenly curved, the narrower part
at about a right angle to the wider base
(Figs. 104, 105, 109, 111, 115, 116), while
that of M. foxi is at about a 45 angle.

Metepeira grandiosa palustris
Chamberlin and Ivie

Figures 97, 98, 101-105; Map 2

Note. Emerton's bog variety of M. Ial)y-
rinfliea was this subspecies, although some
specimens in the collections from Mount
Lincoln, Colorado, considered here to be

grandiosa alpiiia, were also labeled bog
\ariety. The only name available is palus-
tris.

Diagnosis and Variation. Metepeira gran-
diosa palustris is readily recognized by its
black coxae (Fig. 98). It is slightly larger
than M. grandiosa alpina. Total length of
females varies 4.7 to 7.6 mm, carapace 1.9
to 2.9 mm long, 1.6 to 2.2 mm wide. Total
length of males 3.1 to 3.8 mm, carapace 1.7
to 1.9 mm long, 1.4 to 1.5 mm wide. First
patella and tibia of female is the same
length as carapace to 1.2 times its length;
that of males, 1.5 its length.

Natural History. Emerton collected speci-
mens in bogs in Maine; they also have been
collected in a marsh in the Adirondacks, in
low spruce (Picea sp.) in Quebec and in a
field in Wisconsin.

Distrihution. Canada from Nova Scotia
to British Columbia, south to Maine, New
York, South Dakota (Map 2).

Metepeira grandiosa alpina
Chamberhn and Ivie

Figures 99, 100, 106-111; Map 2

Note. The types of names M. dakota and
M. alpina belong to this subspecies. Al-
though the name M. dakota has page pri-
ority, as first revisor 1 chose M. alpina
since the type locality is in the middle of
the range of the subspecies and is a place
where it is common, while Noonan, North
Dakota, the type locality of M. dakota, is at
the border of the range.

Diagnosis and Variation. This includes
the smallest sizes and the most abundant
populations. The scape of the epigynum is
wide at the base (Fig. 106) and the palpal
embolus is int(>rmediate in length (Fig.
111). The coxae are ne\'er black, but inter-
mediates are found among the eastern-most
specimens (see M. grandiosa palustris).
Total length of females is from 4.0 to 6.8

Figures 101-105. Metepeira grandiosa palustris Chamberlin and Ivie (Nova Scotia). 101-103. Epigynum:
101. Ventral. 102. Posterior. 103. Posterior, cleared. 104. Left male palpus, mesal. 105. Median apophysis
and embolus.

Metepeira, Kaira, Aculepeira • Levi 213

Figures 106-111. M. grandiosa alpina Chamberlin and Ivie (northern Arizona). 106-108. Epigynum: 106.
Ventral. 107. Posterior. 108. Posterior, cleared. 109. Male palpus, mesal. 110. Palpus, ventral. 111. Median
apophysis and embolus.

Figures 112-116. M. grandiosa grandiosa Chamberlin and Ivie (Riverside Co., California). 112-114. Epigynum:
112. Ventral. 113. Posterior. 114. Posterior, cleared. 115. Male palpus, mesal. 116. Median apophysis and
embolus.

Scale lines. 0.1 mm.

214 Bulletin Musruiu of Compaiativc Zoolugy, Vol 148, No. 5

mm, carapace 1.9 to 2.9 mm long, 1.5 to
2.2 mm wide. Total length of mak>s ranges
from 3.1 to 5.3 mm, carapace 1.8 to 2.4 mm
long, 1.4 to 1.8 mm wide. The first patella
and tibia is 1.0 to 1.5 times the length of the
carapace in females, 1.5 to 1.7 times the
length in males.

Natural History. Adult males have been
collected in June and July, females from
June to August. Metepeira ^randiosa alpiim
has been collected by sweeping a meadow
in South Dakota, from a meadow , browsed
aspen, bunchgrass {?Sporoh()lus airoides)
and with sage {Artemisia) in Wyoming.
Additional specimens have been found in a
meadow, in oak juniper (two collections),
on sagebrush (Artemisia) from a dry hill-
side in Colorado ( two collections ) and from
sagebrush in Oregon. Specimens are com-
monly collected with M. foxi. Their habitats
must be similar.

Distribution. Rocky Mountain area of
Saskatchewan, North Dakota, Oklahoma, to
British Columbia, Oregon and south to
Chihuahua (Map 2).

Metepeira grandiosa grandiosa
Chamberlin and Ivie

Figures 112-116; Map 2

Note. Types with the names M. grandiosa
and M. palomara belong to this subspecies.

Diagnosis and Variation. Specimens of
this subspecies are more variable than those
of M. g. aJpina but tend to be larger in
size. Often the scape of the female epigy-
num is narrower than that of the other sub-
species; it has a median ridge (Fig. 112)
and the distal part of the embolus is heavier
(Figs. 115-116). Total length of females
varies from 5.4 to 8.5 mm, carapace 2.3 to
3.5 mm long, 1.9 to 3.0 mm wide. Total
length of males is 3.5 to 5.1 mm, carapace
1.9 to 2.7 nun long, 1.4 to 1.9 mm wide.
First patella and tibia of female is 1.3 times
the length of the carapace, that of the male,
1.3 to 1.8 times.

Natural History. Mal(\s have been col-
lected from March to October, females from
April to September. They have been col-

lected in the same area as M. cra.ssipes, in
Riverside County, California. The species
was abundant only in spring, not in fall.

Distribution. British Columbia to Cali-
fornia (Map 2).

Kaira O.P.-Cambridge

Kaim O.P.-Cambridge, 1889, Biologia Centrali-
Ainericana, Araneidea, 1: 56. Type species K.
flihhewsa O.P.-Cambridge, 1889 designated by
F. P. -Cambridge, 1904, Biologia Centrali-Ameri-
cana, Araneidea, 2: 522. The name Kaira is of
feminine gender.

Caira Simon, 1895, Histoire Naturelle des Araignees,
2nd ed., 1: 894. New name for Kaira (pre-
sumably l^ecause the Latin alphabet lacks a K),
an invalid emendation.

Pronarachnc Mello-Leitao, 1937, An. Acad. Bra-
sileira Sci., 11: 9. Type species by monotypy,
P. aries Mello-Leitao, 1937 {?= Kaira alba).
NEW SYNONYMY.

Macpos Mello-Leitao, 1969, Rev. Chilena Hist.
Natur. 43: 59. Type species by monotypy, M.
monstrosus Mello-Leitao. NE^^^ SYNONYMY.

Diagnosis. Kaira differs from Metepeira
and Acidepeira by the shape of the abdo-
men, which is attached in the center of its
anterior side and has the longest axis almost
at a right angle to that of the cephalothorax
(Figs. 123, 125); by the tuberculate dorsal
humps on the abdomen of females (Figs.
122, 134, 144, 146); by the heavily armed
distal portion of the legs of females that
have short metatarsi and tarsi (Figs. 122,
144, 146); and by lacking the black pigmen-
tation around the median white mark on the
venter of the abdomen (Fig. 123).

Coloration. Tlie color of all species ap-
pears yellowish white, with only scattered
black pigment (Figs. 122, 126, 144-146);
there is a central white mark on the venter
of the abdomen. The males are darker than
the females (Fig. 126). It is possible that
living specimens are green.

Description. Carapace noticeably narrow
in the head region (Figs. 121, 122, 126).
Chelicerae armed with three long teeth on
the anterior margin (one of the three
slijihtlv shorter than the others) and with
two denticles on the posterior margin (Fig.
121). Endites short, unlike those of Mete-

Metepeira, Kaira, Aculepeira • Le

215

peira. Height of the clypeiis ecjiials that of
the anterior median eyes or sHglitly higher
(Fig. 121). Abdomen higher than long, has
tubercuhite dorsal humps and covered with
short setae (Figs. 122, 12.3). Tibiae slightly
sinuous; the metatarsi and tarsi noticeably
short and curved (Fig. 123). Distal part of
the legs hea\ily armed, in K. (ili)a w ith short
macrosetae (Fig. 121) and in K. salrino with
both short and long macrosetae (Figs. 144,
146).

Males are smaller than females. Palpal
femiu" lacks the proximal tooth; palpal pa-
tella lacks macrosetae. Neither coxae nor
legs of males modified; and they lack macro-
setae present in the female. Male's ab-
dominal humps much smaller than female's
and mav not be tuberculate ( Figs. 126,
135, 138).

Genitalia. The genitalia are surprisingly
similar to those of Metepeira. The epigy-
num is small and as weakly sclerotized as
that of Metepeira (Figs. il7, 130, 141);
the openings are on each side of the pos-
terior face leading into pockets that funnel
into the large spherical seminal receptacles
(Figs. 120, 123, 143). The palpus has a
strongly sclerotized median apophysis wdih
a distal row of teeth and two flagella ( Figs.
127-129, 136, 137, 139, 140) siniilar to that
of some species of Metepeira. Tlie embolus
(e in Fig. 129), as in Metepeira, has a lobe
below the duct-bearing portion. A long
terminal apophysis (a) is separated from
the embolus by a distal hematodocha (dh
in Fig. 129). It is not known if the virgin
embolus has a cap that is transferred to the
female when mating.

Natural History. Nothing is known of
habits; the species are so rare that one can
only speculate what the habits might be.
Do the armed distal articles of the legs
(Figs. 122, 123) indicate that the spider
does not make a web, but instead catches
insects wdth the legs in a crabspider fashion,
as do some other Araneidae?

Species. There are so few specimens in
collections that it is difficult to decide the
limits of species and to match males and

females. Species appear to differ by the
shape of the abdomen (Figs. 124, 134, 145),
by the proportions of the scape of the
epigynum (Figs. 118, 131, 141) and by the
shape of the conductor of the palpus (c in
Fig. 129, Figs. 128, 137, 140) (in contrast
to many species of Metepeira where the
conductors resemble each other ) . It is prob-
ably not safe to identify specimens by ab-
dominal humps alone. The humps of two
females of K. .sa])ino are very different. The
reason for this is that one has probaljly just
molted (Figs. 146, 147), while the other
has very recentlv produced eggs ( Figs. 144,
145).

Relationships. The genitalia of Kaira re-
semble Metepeira to such an extent that a
common ancestor should be assumed. An-
other similarity is the white spot on the
venter of the abdomen. Kaira shares the
closest relationship with the M. foxi group,
which also has teeth on the distal keel of the
median apophysis (m in Fig. 129). I con-
sidered the M. foxi group the least special-
ized of Metepeira.

Distribution. Kaira species are known
only from the wanner and tropical parts of
the Americas.

Key to fetiiale Kaira north of Mexico

1. Epigynum with a longitudinal, longer than
wide, transparent keel or hook (Figs. 117,
130 ) ^ 2

- Epigynum with a transverse, wider than
long, transparent scape (Fig. 141); Ari-
zona (Map 3) .subino

2(1) Epigynum having a hook-shaped scape
(Fig. 132); southern Texas to South Amer-
ica (Map 3) alticcntcr

- Epigynum having a keel-shaped scape, of
variable profile (Fig. 119): Xortli Carolina
and Missouri to northeastern Mexico ( Map

3 ) alba

Key to male FLura north of Mexico

1. Palpus with a light swelling on distal end
of conductor, conductor without proximal
pocket (Figs. 128, 129, 137); terminal
apophvsis sharplv pointed ( Figs. 127, 129,
136 ) ' 2

- Conductor with a dark distal swelling over-
hanging suhdi\ided lateral pockets (Fig.
140); terminal apophysis bluntly pointed

216 BuUcliii Mti.sctiiii of Coniparatwc Zoology, Vol 148, No. 5

Map 3. Distribution of Kaira species found north of Mexico. Circles, females; open circles, juvenile specimens;
squares, males; open squares, juvenile males.

( Fig. 139 ) ; ? South Carolina, Arkansas,
northern Texas (Map 3) hitcac

2(1) Proximal flagelluni of median apophysis
longer than distal one (Figs. 127, 128);
tip of conduetor pointed mesally, toward
median apophNsis (Fig. 128); North Caro-
lina and Missouri to northeastehi Mexico
(Map 3) aiha

- Flagella of median apophysis equal in
length (Figs. 136, 137); tip of conductor
pointed laterally, away from median apo-
physis (Fig. 137); Texas to South America
(Map 3) altiventei

Kaira alba (Hentz)

Figures 117-129; Map 3

Kjicira alba Hentz, 1850, J. Boston Soc. Natur.
Hist., 6: 20, pi. 3, fig. 7. Female from North
Carolina in tlie Boston Natural History Society,
destroyed.

Kaira alba: — Keyserhng, 1892, Spinnen Amerikas,

4: 64, pi. 3, fig. 50, $. McCook, 1893, Ameri-
can Spiders, 3: 202, pi. 13, fig. 3, $. Comstock,
1912, Spider Book, p. 450, fig. 461, 9 ; 1940,
Spider Book, rev. ed., p. 464, fig. 461, ?.
Roewer, 1942, Katalog der Araneae, 1: 904.
Caiia alba: — Bonnet, 1956, Bibliographia Araneo-
rum, 2: 924.

Description. Female from Florida. Cara-
pace yellow-white with a median darker
line of black spots and some straggly white
setae (Fig. 122). Sternum with tiny black
spots. Legs yellowish white with tiny black
spots and narrow broken black bands on
venter (Fig. 123). Dorsum of abdomen
with a black mark between humps, sides
with tiny black spots (Figs. 122, 124, 126).
Venter spotted black. Posterior median
eyes subequal to anterior medians; laterals
0.8 diameter of anterior median eyes. An-

Metepeira, Kaira, Aculepeira • Levi 217

117

119

Figures 117-129. Kaira alba (Hentz). 117-124. Female: 117-120. Epigynum: 117. Ventral. 118. Posterior. 119.
Lateral. 120. Posterior, cleared. 121. Eye region and chelicerae. 122. Female, dorsal. 123. Female, lateral. 124.
Abdomen, posterior. 125-129. Male; 125. Lateral. 126. Dorsal. 127-129. Left palpus: 127. Mesal. 128. Ventral.
129. Expanded, subventral.

Abbreviations, a, terminal apophysis; c, conductor: dh, distal hematodocha; e, embolus; h, hematodocha; m,
median apophysis; r, radix; t, tegulum.

Scale lines. 0.1 mm except Figs. 122-126, 1.0 mm.

218 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

terior median eyes 1.5 diameters apart, 2.5
from laterals. Posterior median eyes 1.4
diameters apart. Height of clypeus equals
diameter of anterior median eyes. Anterior
margin of chelicerae has two long teeth
distally and two smaller ones proximally.
On the posterior margin are two distal teeth
and two small proximal denticles. Abdomen
much higher than long \\'ith tuberculate
humps (Figs. 123, 124). Total length 6.5
mm. Carapace 2.9 mm long, 2.7 mm wide.
First femur, 3.6 mm; patella and tibia, 4.4
mm; metatarsus, 2.3 mm; tarsus, 1.1 mm.
Second patella and tibia, 3.6 mm; third, 2.2
mm; fourth, 2.8 nun.

Male coloration darker than that of fe-
male (Fig. 126). Anterior median eyes
slightly more than their diameter apart,
slightly less than one from laterals. Posterior
median eyes slightly more than their diam-
eter apart, 1.5 from laterals. Abdomen ver-
tical with large humps that are slightly ir-
regular, only faintly indicating tubercles
(Fig. 126). Total length 3.0 mm. Carapace
1.4 mm long, 1.1 mm wide. First femur,
1.7 mm; patella and tibia, 2.1 mm; meta-
tarsus, 1.1 mm; tarsus, 0.7 mm. Second
patella and tibia, 1.6 mm; third, 0.9 mm;
fourth, 1.2 mm.

Variation. The shape of the two humps
of the abdomen is variable. Females vary
in total length from 6.5 to 7.3 mm, carapace
2.9 to 3.1 mm long; males from 2.6 to 3.0
mm in total length.

Dia<i,nosis-. The median longitudinal keel
of the lightly sclerotized epigynum (Figs.
117, 119) separates the species from K.
altiventer. The shape of the conductor of
the palpus (Figs. 128, 129) and the pointed
terminal apophysis (Figs. 127, 129) sepa-
rates the species from K. hiteae.

Natural History. Females have been col-
lected in all seasons, males only in early
summtM-. In Florida females have been col-
lected on grass stems; in mangrove ham-
mocks, and in shrubs bordering a red and
white mangrove hammock; a male from
Arkansas came from oak-hickory bmsh. We
do not know the web of this uncommon

species. Males are collected by sweeping
vegetation.

Distribution. From North Carolina, Ten-
nessee, Arkansas to northeastern Mexico
(Map 3). Juvenile specimens have been
mapped with open circles.

Records. NortJi Carolitui. Guilford Co.:
5.2 mi. north of Greensboro, 10 June 1953,
6 (R.D. Barnes). Tennessee. Washington
Co.: Johnson City, 12 June 1951, $ (O.
Bryant). Florida. Jackson Co.: 12 May
1935, juv. (H.K. Wallace). Alachua Co.:
Gainesville, 10 Oct. 1963, 2 (L.A. Hetrick).
Lake Co.: Leesburg, March 1954, $ (M.
Statham). Seminole Co.: Sanford, Sept.
1927, juv^ (Stone). Osceola Co.: Kissim-
mee, juv. (N. Banks). Sarasota Co.: Engle-
wood, 1 Apr. 1938, juv. (W.J. Gertsch).
Dade Co.: Matheson Hammock, 20 June
1964, 9 (K. Stone). Monroe Co.: 2 mi.
north of Flamingo, 21 June 1964, 9 (K.
Stone). Mississippi. Wilkinson Co.: Cen-
treville, 1944, S (A.F. Archer). Missouri.
Stoddard Co.: Ardeola, 22 July 1950, 9
(H. Exline). Arkansas. Washington Co.:
15 mi. west of Prairie Grove, S (M. Hite).
Randolph Co.: 16 June 1963, $ (Exline
Coll.). Texas. Denton Co.?: Decator, 1945,
c^ (H. Exline). Hidalgo Co.: Edinburg,
Sept.-Dec. 1933, 9 (S. Mulaik). Mexico.
Tamaulipas. Soto La Marina, 16 May 1952,
9 (W.J. Gertsch). San Luis Potosi. Valles,
July 1959, juv. (L. Steude).

Kaira altiventer O.P. -Cambridge
Figures 130-137; IVlap 3

Kaira altiventer O.P.-Cambridge, 1889, Biologia
Centrali-Americana, Araneidea, 1: 56, pi. 3, fig.
13, 9 . Female holotype from Veragua, Panama,
in the British Museum, Natural History, ex-
amined. Kevserling, 1892, Spinnen Amerikas,
4: 62, pi. 3, fig. 48, 9. F.O.P.-Cambridge, 1904,
Biologia Centrah-Americana, Araneidea, 2: 522,
pi. 51, fig. 10, 9 . NEW SYNONYMY.

? Kaira obtusa Kevserling, 1892, Spinnen Amerikas,
4: 66, pi. 3, tig. 51, juvenile holotype from
Taguara, Rio Grande do Sul, examined.

Caira spinosa Simon, 1896, Ann. Soc. Entomol.
France, 65: 478. Female lectotype here desig-
nated and juvenile paralectotype from Sao Paulo
de Oli\en^a, Amazonica, Brazil and Pebas, Peru

Metepeira, Kaira, Aculepeira ' Levi 219

Figures 130-137. Kaira altiventer O. P. -Cambridge. 130-134. Female: 130-133. Epigynum: 130. Ventral.
131. Posterior. 132. Lateral. 133. Posterior, cleared. 134. Abdomen, posterior. 135-137. Male: 135. Dorsal.
136, 137. Left palpus: 136. Mesal. 137. Ventral.

Figures 138-140. K. hiteae. male. 138. Dorsal. 139, 140. Palpus: 139. 140. Palpus: 139. Mesal. 140. Ventral.

Scale lines. 0.1 mm except Figs. 134, 135, 138, 1.0 mm.

in the Museum National d'Histoire Naturelle, Museu Nacional, Rio de Taneiro, examined.

Paris, examined. NEW SYNONYMY. NEW SYNONYMY.
Pronarachne aries Mello-Leitao, 1937, An. Acad.

Brasileira Sci., 9: 9, fig. 10, $. Female holo- Description. Female from Texas. Cara-

type from Itatiaia, Rio Grande do Sul in the pace \ ellow-white with some tiny orange

220 Bulletin Museum of Coinparaticc Zoology, Vol. 148, No. 5

dots and some white pigment streaks. Dor-
sum of abdomen \\ ith some tiny black and
orange spots. Venter with spots. Legs yel-
low-white with tiny orange spots. Eyes
subequal in size. Anterior median eyes 1.2
diameters apart, 2.5 from laterals. Posterior
median eyes one diameter apart. Height of
the clypeus equals about the diameter of
the anterior median eyes. Total length 6.0
mm. Carapace 2.9 mm long, 2.3 mm wide.
First femur, 3.0 mm; patella and tibia, 3.9
mm; metatarsus, 2.0 mm; tarsus, 0.9 mm.
Second patella and tibia, 3.1 mm; third, 1.6
mm; fourth, 2.3 mm.

Male. Abdomen shield-shaped, humps
dorsally directed (Fig. 135). Total length
2.0 mm. Carapace 1.2 mm long, 1.0 mm
wide. First femur, 1.1 mm; patella and
tibia, 1.3 mm; metatarsus, 0.8 mm; tarsus,
0.4 mm. Second patella and tibia, 1.0 mm;
tliird, 0.6 mm; fourth, 0.8 mm.

Note. It is not certain that the males and
females described and ilhistrated belong
together.

Variation. The palpi of the two males
differ slightly; the illustrations were made
of the male from El Salvador (Figs. 136,
137). The specimen from Costa Rica dif-
fers in that the two flagella of the median
apophysis are more straight and the con-
ductor seems slightly turned on its longi-
tudinal axis facing the median apophysis,
appearing less pointed.

Diagnosis. Females can be separated
from Kaira alba by the hook-shaped scape
of tlic cpigynum ( Fig. 132 ) . Males can be
separated from those of K. alba, by the di-
rection of the distal swelling of the con-
ductor (Fig. 137), and by the absence of
a lateral fold (Fig. 137) from K. hiteae.

Di.stri])ution. Texas to South America
(Map 3).

Records. Texas. Hidalgo Co.: Edinburg,
6 Dec. 1935, 9 (S. Mulaik). EJ Salvador.
San Salvador, Jan.-March, 1954, 6 (J.B.
Boursot). Costa Rica. Cuanacaste Prov.:
4 km NW Canas La Pacifica, 7 Feb.-2
March 1975, i (R. Coville from Trijpar-
g,ilinn nitiduin muddauber nest). Canal

Zone. Ft. Sherman, Aug. 1939, juv. (A.M.
Chickering).

Kaira hiteae new species
Figures 138-140; Map 3

Holotype. Male from Cove Creek Valley,
9.3 km west of Prairie Grove, Boston Moun-
tains, 300 m elevation, Washington County,
Arkansas, 26 July to 7 August 1956 ( Maxine
Hite ) in the Museum of Comparative Zool-
ogy. The species is named after M. Hite,
the collector of several specimens of these
rare species.

Description. Male. Carapace yellowish,
lighter in center. Legs yellowish, very in-
distinctly banded. Abdomen with indis-
tinct transverse bands, venter white. Eyes
subequal in size, except anterior laterals
0.8 diameter of anterior median eyes. An-
terior median eyes 1.3 diameter apart, 1.3
from laterals. Posterior median eyes slightly
less than their diameter apart, slightly more
than two diameters from laterals. Height
of the clypeus equals diameter of the an-
terior median eyes. Endites are short and
s(iuare. None of the legs have strong
macrosetae. Abdomen is shield-shaped
(Fig. 138). Total length 2.6 mm. Carapace
1.3 mm long, 1.2 mm wide. First femur,
1.5 mm; patella and tibia, 1.9 mm; meta-
tarsus, 1.2 mm; tarsus, 0.7 mm. Second
patella and tibia, 1.4 mm; third, 0.8 mm;
fourth, 1.1 mm.

Note. At first it appeared that these
males Ix'longed with the females of K.
sabino. but I now doubt this.

Diaiinosis. Kaira hiteae differ from K.
alba Iw lacking black pigment in the center
of the carapace and between the humps
( Fig. 138); in K. hiteae this area is Hghtest.
They differ too by having transverse bands
on the abdomen and on the single point of
the humps (Fig. 1.38). Tlie males have a
more blunt terminal apophysis (Fig. 139),
the conductor has an edge laterally (Fig.
140) and a slightly more distal attachment
of the flagcllae to the median apophysis
(Figs. 139, 140).

Natural History. One male from Dallas

Metepeira, Kaira, AcuLEPEiRA • Lcvi 221

145

147

Figures 141-147. Kaira sabino n. sp., female. 141-143. Epigynum: 141. Ventral. 142. Posterior. 143. Posterior,
cleared. 144-147. Female; 144, 146. Dorsal. 145, 147. Abdomen, posterior. 144, 145. Paratype (Portal, Ari-
zona). 146, 147. Holotype (Sabino, Arizona).

Scale lines. 0.1 mm except Figs. 141-144, 1.0 mm.

County, Texas had a label stating it had
been collected with spiders from "road,
woods, herbs and shrubs."

Distribution. South Carolina, Arkansas,
Texas (Map 3).

Records. South Carolina. York Co.: 3.4
mi. west of Sharon, 4 Aug. 1953, juv. S ( R.
Barnes). Arkansas. Washington County:
Cove Creek Valley, Boston Mts., 1000 ft,
26 July, 7 Aug. 1956, 2 ^ (M. Kite), one
deposited in Canadian National Collection.

Texas. Dallas Co.: Coit, 2 Aug. 1940, S
(S. Jones). Grayson Co.: Sherman, 25 July
1963, 2 c^ (K.W. Haller). (All but first
listed specimen have been marked as para-
types. )

Kaira sabino new species
Figures 141-147; Map 3

Holoytpe. Female from Sabino [? Can-
>'on, Pima County], Arizona, 2 September

222 BuUrtin Museum of Comparative Zoology, Vol. 148, No. 5

1932 (R. Flock) in the Museum of Com-
IDarative Zoology. The specific name is a
noun in apposition after the type locality.

Note. According to J. Beatty (personal
communication), there may be numerous
locaHties with the name of Sabino, Arizona.
W.C. Barnes. 1960, Arizomi Place Names
has listed onl>- two, both in Pima County,
one of which, Sabino Canyon, is a common
collecting site.

Description. Female holotype. Carapace
light brown, darker brown on sides. Ster-
num darker on sides. Legs with indistinct
narrow bands. Dorsum of abdomen with
transverse bands (Figs. 145, 147). Sides
light, venter with white pigment. Posterior
median eyes 1.5 diameters of anterior me-
dians, anterior laterals 1.2; posterior laterals
1.5 diameters of anterior medians. Anterior
median eyes 2 diameters apart, 2.5 diam-
eters from laterals. Posterior median eyes
their diameter apart. Height of the clypeus
equals 1.5 diameters of the anterior median
eyes. First metatarsus curved and armed
with strong and weak setae (Figs. 144, 146).
Abdomen appears shield-shaped, having
small tubercles on the humps (the abdo-
men, however, is in very poor condition).
Total k>ngth 5.8 mm. Carapace 2.4 mm
long, 1.9 mm wide. First femur, 3.2 mm;
patella and tibia, 4.0 mm; metatarsus, 2.3
mm; tarsus, 1.0 mm. Second patella and
tibia, 3.1 mm; third, 1.7 mm; fourth, 2.2 mm.

Note. It appeared at first that these fe-
males belonged with the males of K. hiteae,
but I now doubt this.

Variation. The second specimen, although
having a similar epigynum to the first and
doubtlessly of the same species, is quite
different in appearance: it has scattered
black pigment and many more abdominal
tubercles (Figs. 144, 145). Since a fluffy
egg-sac is in the vial, it must have oviposited
and the abdomen shriveled.

Diagnosis. The wide short scape of the
epigynum, which appears to be a transverse
lip ( Fig. 141 ) , is very distinct from the
narrow keel or hook of the two other species.

Paratype. Arizona. Cochise Co.: 1 mi.

east, 7 mi. north of Portal on San Simon
Road, 17 July 197.3, 5 (A. Jung).

Aculepeira Chamberlin and Ivie

Aculepeira Chamberlin and Ivie, 1942, Bull. Univ.
Utah, biol. sen, 7(1): 75. Type species desig-
nated l)y original authors as Epcira aculcata
Emerton, which these authors thought to be the
name of A. carbonarioides; actually E. aculeata
is a synonym of A. packardi (Thorell). The
name Aculepeira is feminine.

Diagnosis. Aculepeira, like Metepeira, has
a median, ventral white streak on the abdo-
men but differs in both sexes by the elon-
gate, egg-shaped abdomen, in the female
by the large, sclerotized epigynum with a
large scape (Figs. 148, 162), and in the
male by the large palpus with long, prom-
inent, median apophysis with two flagella
(Fis. 159, 161) and a large boat-shaped to
disk-shaped conductor ( Figs. 158, 160, 170,
172). Aculepeira differs from the related
Araneus by having an elongate, egg-shaped
abdomen that is widest anteriorly (Figs.
154, 166 ) ; the abdomen of the large Araneus
usually ranges in shape from spherical to
longer than wide and may have humps.
UnUke any of the large Araneus species,
Aculepeira species have a median, ventral
white mark on the abdomen (Figs. 155,
168).

Description. Carapace with eye area and
thoracic rim light and thorax dark covered
by white down; sternum, brownish black.
Legs banded; abdomen with a dorsal longi-
tudinal lobed band (Figs. 154, 166) and
a ventral white mark on black ( Figs. 155,
168). Eyes subecjual in size, laterals some-
times smaller. Anterior median eyes slightly
more than their diameter apart, posterior
median eyes their diameter apart; in female,
slightly closer spaced than in male. Height
of the clypeus, about 1.5 to almost 2 diam-
eters of the anterior median eyes (Fig. 174).
The thorax with a shallow depression ( Figs.
154, 166, 174). Abdomen, chicken-egg-
shaped, longer than wide, widest anteriorly
(Figs. 154, 166) and covered with setae;
very hairy in A. carhonaria and A. car-
bonarioides.

Metepeira, Kaira, Aculepeira • Levi 22S

Plate 6. Aculepeira packardi. Above, spider in web; below, spider in a disc-shaped retreat in vegetation.

Male smaller than female (Fig. 157),
chelicerae small and anteriorly concave.
Endites with a tooth laterally facing an-
other tooth at the proximal end of palpal
femur. First coxae with a hook fitting into
of second femur. The second

groove

coxae with a median ventral pointed hump
in most species and the fourth trochanter
with a posterior macroseta in most species
( Fig. 175 ) . Second tibia distally wider with

short macrosetae, one of which is on a pro-
jecting hump (Figs. 156, 159).

Genitalia similar to those of Araneus,
especially to A. washingtoni Levi. Epigy-
num, an annulate scape on a base (Figs.
148-152, 162-163) and lamellae on each
side posterodorsally (Figs. 150, 164). Pal-
pal patella with two macrosetae (Fig. 172);
palpal parts like those of A. ivashingtoni,
except for a long median apophysis with

224 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Plate 7. Aculepeira carbonarioides. Top, female; middle and bottom, orb-web in talus in Colorado. Webs
dusted with cornstarch. Viscid area of web approximately 15-20 cm diameter.

Metepeira, Kaira, AcuLEPEiRA • Levi 225

two proximal flagclla (m in Figs. 161, 173).
Conductor a deep sclerotized boat-shaped
to disk-shaped stiaicture (Figs. 158, 160,
170, 172); embolus (e in Figs. 161, 173)
hidden in contracted palpus by the terminal
apophysis (a in Figs. 161, 173) and lacking
a cap in virgin males. Terminal apophysis
a long sclerotized prong with a hook at its
base (a in Figs. 161, 173). Subterminal
apophysis acutely pointed in A. packardi,
more blunt in A. carhonarioides but hidden
underneath the terminal apophysis (Figs.
158, 170 and sa in Figs. 161, 173).

Eurasian Species. The three common
European species, frequently confused with
the North American ones, have been
illustrated (Figs. 187-217). I had first
planned to revise Aculepeira worldwide,
but lack of specimens, especially males
from the Near East, made this impossible.
There were nomenclatural and biological
problems that I could not resolve (see
Methods above).

Thorell (1870) gave the name Epeira
victoria to illustrations by C. L. Koch of
a spider from Germany and France on low
plants in meadows and the border of forests
(C. L. Koch, 1834, 1839). Drensky (1943)
thought A. victoria to be the same as A.
armida, although the latter species is not
known to occur in Germany.

The specimens from Switzerland labeled
A. victoria in the British Museum and the
Naturhistorisches Museum, Basel are prob-
ably A. armida. But there are female speci-
mens from Dalmatia and Samarkand, USSR,
in the Natural History Museum, Stockholm
that may be distinct ( Figs. 226-231 ). These
have a shorter scape and wide lamellae
posterodorsal to the epigynum.

Epeira armida was described from south-
ern France and northern Israel. Are the
populations found in these two areas the
same species? I examined several collec-
tions from Turkey and all but two were
females. None looked quite ahke. Karol
( 1964, 1966 ) recently described two species
(A. vachoni, A. karapagi) from females

from Turkey without making comparative
reference to the other species reported from
th(^ area.

The type of A. vachoni was examined
and it appeared to be a female A. ceropegia
having the scape of the epigynum torn off.
Aculepeira karapagi appears to be a dis-
tinct species previously described as Epeira
similis Nosek, 1905 (a name preoccupied)
and subsequently renamed E. noseki Strand,
1907.

Natural History. Both American species
make fairly large orbs and one is found in
extreme habitats not otherwise frequented
by large orb-weavers. Aculepeira packardi
has a retreat (Plate 6), A. carhonarioides
does not (Plate 7). Aculepeira packardi
males are more commonly collected than
females, usually by sweeping, but A. car-
honarioides males are rare in collections,
presumably because it is difficult to collect
among boulders.

Note on Names. Both Keyserling and
Emerton realized there are two species in
North America, but each called a different
one Epeira carhonaria and gave the second
a new name. Neither American species is
A. carhonaria.

Key to American Aculepeira species

1. Dorsum of abdomen in l)oth sexes with
median, paired, upside-down, connna-sliaped
marks or posteriorly directed lobes ( Figs.
154, 157); width of scape of epigynum more
than half the width of base (Fig. 148) or
scape torn off (Fig. 152); epigynum in pos-
terior view with median area framed by
convexly lobed lateral pieces ( Figs. 150,
151); conductor of palpus relatively wide
with ventral lip (Figs. 160, 176-181) _....,
packardi

- Dorsum of abdomen with paired white pig-
ment patches, but no lateral, posteriorly di-
rected lobes (Fig. 166), or with indistinct
dark pattern of paired spots (Fig. 167);
width of scape of epigynum less than a third
the width of base (Figs. 162, 182, 185);
scape rarely torn off; epigynum in posterior
\iew with median area framed by concavely
curved lateral pieces (Figs. 164, 165, 183,
186); conductor of palpus narrow without
a lip around ventral margin ( Figs. 170, 172 )
carhonarioides

226 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Aculepeira carbonarioides

Map. 4. Distribution of North American Aculepeira species.

Metepeira, Kaira, Aculepeira • Levi 227

Figures 148-161. Aculepeira packardi (Thorell). 148-153. Epigynum: 148, 152, 153. Ventral. 149. Lateral. 150.
Posterior. 151. Posterior, cleared. 152. Scape torn off. 148-151. (California). 152. (Nevada). 153. (Colorado).
154. Female, dorsal. 155. Female abdomen and sternum, ventral. 156. Left male second patella and tibia,
dorsal. 157. Male, dorsal. 158-161. Left male palpus: 158. Apical. 159. Mesal. 160. Ventral. 161. Expanded,
ij mesal view.

Abbreviations, a, terminal apophysis; c, conductor; dh, distal hematodocfia; e, embolus; i, stipes; m, median
apophysis; r, radix; sa, subterminal apophysis.

Scale lines. 1.0 mm.

228 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

Aculepeira packardi (Thorell)

Plate 6; Figures 148-161, 174, 176-181;
Map 4

Epcira })ackardii Thorell, 1875, Proc. Boston Soc.
Natur. Hist., 17: 490. Left palpus of holotype
from Square Island, Labrador in tiie Natural
History Museum, Stockliolm, examined; the re-
maining parts of specimen probably destroyed.

Epcira acideata Enierton, 1877, Bull. U.S. Geol.
Surv. Territ., 3: 528, fig. 18, 9. Female holo-
type from Gray's Peak [Front Range, Summit
Co.], Colorado, lost. Emerton, 1894, Trans.
Connecticut Acad. Sci., 9: 405, pi. 1, fig. 4 a,
c, e, $, $. NEW SYNONYMY.

Epeira carhonaria: — Keyserling, 1892, Spinnen
Amerikas, 4: 204, fig. 151, $, $. McCook,
1894, American Spiders, 3: 157, pi. 5, figs. 1,
2, 9 , $ • Misidentification, not A. carhonaria
(L. Koch).

Arancus septentrionalis Kulczynski, 1908, Zap.
Imp. Akad. Naouk, (8) 18: 47, fig. 57, 9.
Female holotype from Sataghai, Adytscha River,
in central Siberia, in Polish Academy of Sciences,
Warsaw, examined. NEW SYNONYMY.

Aculepeira verae Chamberlin and Ivie, 1942, Bull.
Univ. Utah, biol. ser., 7(1): 75, figs. 215, 216,
S . Male holotype from Bear Valley, Idaho in
tlie American Museum of Natural History, ex-
amined. NEW SYNONYMY.

Note. This species has been called cor-
honarius by Gertsch and Ivie in collections
and verae by Chamljcrlin and Ivie.

Description. Female from California:
carapace brown, head region lighter and
covered with white down. Sides of thorax
lighter. Dorsum of abdomen with charac-
teristic pattern (Fig. 154). Total length
11.5 mm. Carapace 4.7 mm long, 3.7 mm
wide. First femur, 4.7 mm; patella and
tibia, 5.S mm; metatarsus, 4.1 mm; tarsus,
1.4 mm. Second patella and tibia, 5.2 mm;
third, 3.1 mm; fourth, 5.0 mm.

Male from California: coloration like fe-
male, but legs less distinctly banded (Fig.
157). Chelicerae with three teeth on an-
terior margin and a small denticle most
distant from base of fang, posterior margin
with tlu-ee teeth. Total length 9.4 mm.
Carapace 4.4 mm long, 3.6 mm wide. First
femur, 5.5 mm; patella and tibia, 6.8 mm;
metatarsus, 4.6 mm; tarsus, 1.4 mm. Second
patella and tibia, 5.3 nun; third, 3.2 nun;
fourth, 5.1 mm.

Variation. The only specimens having a
similar shaped scape on the epigynum or
similarly shaped conductor on the palpus
are those collected together. There is sur-
prising variation in these structures. Speci-
mens with the widest palpal conductor are
sympatric with A. carhonarioides; those
with a narrower conductor are found out-
side the latter species range, providing an
example of character displacement (Figs.
176-181). Total length of females, 5.6 mm
(Yukon) to 16.5 mm (Utah), carapace 2.7
to 6.3 mm long, 2.0 to 5.2 mm wide. Total
length of males 5.0 (Labrador) to 8.9 mm
(Wyoming), carapace 2.6 to 4.4 mm long,
2.1 to 3.7 mm wide.

Diagnosis. This species can be confused
with Neoscona oaxacensis (Keyserling) be-
cause of similar abdomen shape and dorsal
pattern. Neoscona, however, has two pairs
of ventral white patches, while Aculepeira
has a median, ventral white streak on black
(Fig. 155). The dorsal abdominal band,
with posteriorly directed lobes (Figs. 154,
157), separates specimens from A. car- |
honarioides. The scape of the epigynum,
which is usually torn off after mating, is ,
wider than half the width of the base ( Figs.
148, 152) and the conductor is about twice
as long as wide with a ventral lip (Figs.
158, 160, 176-181). The scape of A. car-
])onarioides is less than a third of the width
of the base and the conductor is almost
three times as long as wide and lacks the
ventral lip. The species is close to the Euro-
pean A. ceropegia; but A. ceropegia differs
by the triangular scape, by the median
sclerotized lobe of the base of the epigynum
(Figs. 188-192) and by the enormous, al-
most circular, conductor (Fig. 197).

Natural History. I have collected and
observed this species a nimiber of times in
western Colorado where its webs are found
among the vegetation of lush meadows at
2500 to 3000 m elevation. It is never abun-
dant. Even an entire summer of collecting
may yield only a few specimens. The fe-
males are mature in August in Colorado.
The adult females make a new web ever)'

Metepeira, Kaira, Acvlepeira • Levi 229

Figures 162-173. Aculepeira carbonarioides (Keyseriing), 162-165. Epigynum: 162. Ventral. 163. Lateral.
164. Posterior. 165. Posterior, cleared. 166-168. Female: 166. Dorsal. 167. Abdomen, dorsal. 168. Abdomen
and sternum, ventral. 169-173. Male: 169. Left second patella and tibia, dorsal. 170-173. Left palpus: 170.
Apical. 171. Mesal. 172. Ventral. 173. Submesal, expanded.

Figure 174. A. packardi (Thorell). Female eye region and chelicerae.

Figure 175. A. ceropegia (Walckenaer). Left coxae of male, ventral.

Abbreviations, a, terminal apophysis; c, conductor; e, embolus; h, hematodocha; i, stipes; m, median apoph-
ysis; r, radix; sa, subterminal apophysis; y, cymbium.

Scale lines. 0.1 mm.

230 BuUctiu Museum of Comparative Zoology, Vol. 148, No. 5

night, placing the viscid threads as late as
23:00 h. Once, four spokes from the pre-
vious web were kept. At daytime the female
is found at the side of thc> web in a shallow
retreat ( Plate 6 ) fashioned from several in-
clined leaves tied together and covered
with silk threads. She will actively remove
insects caught during the daytime. A large
bumblebee, however, was left alone. On
12 August 1957 a female was observed eat-
ing a male of the species. Besides lush
meadows in the mountains, the species is
occasionally collected by sweeping grasses
between sagebrush plants (Artemisia).
The temperatures on clear days fluctuated
between 2" C in the morning to 28° C at
noon at one collecting site in Gothic, Colo-
rado (near Crested Butte). A web had 19
radii, with about 25 viscid threads below the
hub and 24 above. The species has been
collected under white spruce ( Picea filauca )
in British Columbia, in sagebrush (Arte-
misia) in New Mexico, and in chaparral
in California. It is found at 1400 to 2700 m
elevations in Utah. During spring in the
southern part of the range, adults of both
sexes may be found. Males are as abundant
in collections as females. They are prob-
aljlv collected by sweeping.

DistriJmtion. Siberia, Alaska to Labrador
and Chihuahua (Map 4). The easternmost
record is CartwTight, Labrador, 7 July 1955,
i (E.E. Sterns, CNC). Another record is
from Pennsylvania, Luzerne Co., Coyning-
ham, $, 17 May 192.3 (W.G. Dietz in the
Academy of Natural Sciences, Philadel-
phia). The specimen was examined in 1968
but could not be located in 1977.

Aculepeira carbonarioides (Keyserling)
Plate 7; Figures 162-173, 182-186;
Map 4

Epcini carhonaria: — Emerton, 1884, Trans. Con-
necticut Acad. Sci., 315: pi. 33, fig. 18, pi. 36,

figs. 18, 19, 9 , $. Emerton, 1894, Trans. Conn.
Acad. Sci., 9: 405, pi. 1, fig. 4b, $ , 4d, $.
Misidcntification, not A. carljonaria (L. Koch).

Epeirci carbonarioides Keyserling, 1892, Spinnen
Anierikas, 4: 206, pi. 10, fig. 152, 9. Female
holotype from Clear Creek, Colorado ( Marx
collection) in the National Museum of Natural
History, Washington, examined. McCook, 1903,
American Spiders, 3: 158, pi. 5, fig. 9, $, $.

? Araneus hi/perboreus Kulczynski, 1908, Zap. imp.
Akad. Naouk, (8) 18: 45, fig. 58, ?. Female
holotype from Dolgulach River, central Siberia,
probably in Polish Academy of Sciences, War-
saw, lost. PROBABLE NEW SYNONYMY.

? Aranea (Epeira) charitonovi Ermolajew, 1928,
Zool. Anz., 77: 209. Specimens from Karagai,
1000 m elcw and Chasinicha River, 3000 m elev.,
Altai Mountains, USSR, deposition unknown.

Araneus vegae Holm, 1970, Entomol. Scandinavica,
1: 198. Male holotype and female allotype from
Nun'amo, St. Lawence Bay [Nunyamo], Siberia
in the Natural History Museum, Stockholm, ex-
amined. NEW SYNONYMY.

Note. There is little doubt that Clear
Creek [County], Colorado (unlike other
Marx localities) is correct. The illustration
Figures 182-184 are made from the types
of A. carbonarioides. Specimens of this spe-
cies from some collections had been labeled
Aranea actileata. Kulczynski's illustration
of A. Jnjper])oreus fits this species. A. cha-
ritonovi differs from A. carhonaria, accord-
ing to the author, by having a narrower
scape; in addition, the illustration fits A.
carhonarioides. Ermolajew considered the
species distinct from A. hijperhoreus be-
cause of larger size; but the size varies
widely in this species.

Description. Female from Colorado:
head region light yellow-brown. Sides of
thorax dark blackish brown with light bor-
der. Dorsum of abdomen very dark, cov-
ered by patches of light pigment and by
tiny white pigment spots, also covered with
lots of setae (Fig. 166). Anterior margin of
chelicerae with four teeth, the second one
from the fang base small; posterior margin

Figures 176-181. Aculepeira packardi (Thorell), variation of conductor of left male palpus, ventral view. 176.
(Kamchatka). 177. (Labrador). 178. (northern British Columbia). 179. (Mohave Desert, California). 180.
(Washington). 181. (Arizona).

Figures 182-186. A. carbonarioides (Keyserling),
184. Lateral. 182-184. (Clear Creek, Colorado).

variation of epigynum. 182, 185. Ventral. 183, 186. Posterior.
185-186. (Grand Teton Mountains, Wyoming).

Metepeira, Kaira, Aculepeira • Levi 231

Figures 187-197. A. ceropegia (Walckenaer) (France, Fig. 191 Switzerland). 187. Left male second patella and
tibia, dorsal. 188-193, Epigynum: 188. Ventral. 189. Ventral, scape torn off. 190-191. Lateral. 192. Pos-
terior. 193. Posterior, cleared. 194. Female, appendages removed. 195. Female abdomen, ventral. 196, 197.
Left male palpus: 196. Mesal. 197. Ventral.

Scale lines. 1.0 mm.

232 Bulletin Miisciiin of Coinparaiitc Zoology, Vol. 148, No. 5

also with four teeth. Total length 12.8 mm.
Carapace 4.(S mm long, 3.9 mm wide. First
femur, 5.3 mm; patella and tibia, 6.3 mm;
metatarsus, 4.7 mm; tarsus, 1.6 mm. Second
patella and tibia, 5.8 mm; third, 3.6 mm;
fourth, 5.3 mm.

Male from Colorado colored like female.
Anterior margin of chelicerae with four
teeth, posterior with three on one side, four
on other. Total length 8.6 mm. Carapace
4.9 mm long, 4.1 mm wide. First femur,
6.3 mm; patella and tibia, 8.4 mm; meta-
tarsus, 6.1 mm; tarsus, 1.5 mm. Second
patella and tibia, 6.6 mm; third, 3.6 mm;
fourth, 5.8 mm.

Varkition. The coxae may be light or
dark or partly dark. The abdominal color
pattern is variable (Figs. 166, 167). Total
length of females 6.2 (New Hampshire) to
15.0 mm (Colorado), carapace 2.6 to 5.4
mm long, 2.0 to 4.5 mm wide. Total length
of males 6.0 (New Hampshire) to 9.1 mm
(Colorado), carapace 3.0 to 4.7 mm long,
2.6 to 3.8 mm wide. Alaska and New
Hampshire specimens are smallest. New
Hampshire males have an embolus that is
shorter and wider, with a base more swollen
and a tip slightly different from specimens
in other localities. On the New Hampshire
specimen's embolus tip, the distal teeth
seem reduced to denticles and there is an
opening of a short filiform portion. In con-
trast, the opening of Rocky Mountain males
is on the top of a short flat portion (e in
Fig. 173). Unfortunately there are no males
from the northern Rocky Mountains, Alaska
or the Arctic in the collections. Perhaps
there are several species. But no consistent
differences were found in epigyna; differ-
ences found (Figs. 182-186) did not vary
with the abdominal color pattern, or other
characters.

Dicifinosls-. Like A. pachirdi, this species
can be confused with Neoscono ooxacensis
(Keyserling), again because of the similar
abdomen shape and dorsal pattern ( Figs.
166, 167). The differences in the white
ventral streak (Fig. 168) separate the two.
However, the abdomen, unlike that of A.

packardi, does not have the lobes of the
median band posteriorly directed (Figs.
166, 167). The scape of the epigynum is
not torn off and is variable in shape but
less than a third the width of the base
(Figs. 162, 182, 185), and the conductor of
the palpus is about three times as long as
wide and lacks a ventral lip (Figs. 170,
172). The similar European A. carhonaria
differs by having a wider scape (Figs. 198,
201 ) and differently shaped conductor of
the palpus (Fig. 207).

Natural History. This species is fairly
common in western Colorado where it
makes its web between boulders of talus
slopes, below and at the timber line (Plate
7). The web forms an angle with the
vertical and the spider rests in the center,
dropping out of the web at the slightest
disturbance. It is then difficult to retrieve
since the creature escapes down among
rocks. After a few minutes, if undisturbed,
it will climb back to the web by means of
a thread it pulled out while falling. I have
not seen the spider sit on stones on the side
of the web. The web sites, 3000 to 3600 m
elevation, probably have extremes of tem-
perature in July from below freezing at
night to up to 30" C during the day.

The species has been collected in Alaska
"between rocks in pika [Ocliototm sp.]
area"; "among rocks at 1900 m" in the
Yukon. The lowest elevation at which it
has been collected in the American Rocky
Mountains was in the Grand Tetons at 2800 •
m elevation. There are few males in col-
lections, reflecting the difficulties of col-
lecting spiders among the boulders.

Distribution. Central Siberia, Alaska to
southern Rocky Mountains (Map 4) (ap-
parently absent from the coast range) in
the east on the Gaspe Peninsula and on
Mount Washington, New Hampshire; both
are areas that escaped glaciation and share
many plant relicts. The easternmost rec-
ord is from Mt. Cartier, Gaspe Peninsula,
Quebec, 5 July 1959, 9 (G. Argus, MCZ)
at 1220 m in spruce-fir. Numerous records

Metepeira, Kaira, Aculepeira • Levi 233

-3a^^^^% 198

Figures 198-207. Aculepeira carbonaria (L. Koch) (Switzerland. Fig. 200, Tyrol). 198-202. Epigynum: 198.
Ventral. 199. Ventral, scape torn off. 200. Lateral. 201. Posterior. 202. Posterior, cleared. 203. Female dorsal,
appendages removed. 204. Female abdomen, ventral. 205. Left second patella and tibia of male, dorsal.
206, 207. Left male palpus: 206. Mesal. 207. Ventral.

Figures 208-216. A. armlda (Audouin) (Spain). 208-211. Epigynum: 208. Ventral. 209. Ventral, scape torn off.
210. Lateral. 211. Posterior. 212. Female, appendages removed. 213. Female abdomen, ventral. 214. Left
second patella and tibia of male, dorsal. 215, 216. Male palpus: 215. Mesal. 216. Ventral.

Scale lines. 1.0 mm.

234 HuUrtin Museum of Comparative Zoology, Vol. 148, No. 5

exist of specimens found between boulders
on Mount Washington.

Aculepeira ceropegia (Walckenaer) new
combination

Figures 175, 187-197

Aranea ceropegia Walckenaer, 1802, Faune Pari-
sienne, 2: 199. Female specimens from Paris,
France, lost. Wiehle, 1931, Tierwelt Deutsch-
lands, 23: 100, figs. 151-155, 9, S- Roewer,
1942, Katalog der Araneae, 1: 783.

Aruncus ceropegius: — Bonnet, 1955, Bibliographia
Araneonmi, 2(1): 454 (European records only).

"> Araneus vachoni Karol, 1964, Bull. Mus. Nat.
d'llist. Natiir., ser. 2, 36: 188. Female holotype
from near Ankara, Turkey, in the Museum
National d'Histoire Naturelle, Paris, examined.
PROBABLE NEW SYNONYMY.

Note. This is a widespread Eurasian
.species that has not been found in North
America. Araneus vachoni appears to be
this species: the torn off base of the scape
is as wide as that of A. ceropegia, the mid-
dle area is sclerotized and the abdominal
pattern is characteristic. Only by finding
a specimen with the scape intact, however,
can the synonymy be determined.

Diagnosis. This species is very similar to
the American A. packardi, having paired,
upside-down, comma-shaped marks on the
abdomen (Fig. 194). It can be separated
from that species by the triangular scape of
the female ej)igynum (which is frequently
torn off ) , by the slightly sclerotized median
area of the epigynum (Figs. 188-192) and
by the very wide, deep conductor of the
male (Fig. 197).

Natural History. According to Wiehle
( 1931 ) A. ceropegia is found in open areas
with the web 50 cm above the groimd; it is
especially abundant in the foothills of the
Alps, the Pyrenees and in the higher moun-
tains to 3000 m. It is known from Scandi-
navia to the Mediterranean area.

Examined Specimens. Sweden. 9 (NRS).
Germany. Bremen, $ (SMF); Hessen:
Eschwege, 9 (H. Homann, SMF); Bavaria:
Obersdorf, $ (SMF). Austria. Tirol. Weis-
senbach, S (H. Wiehle, SMF). Switzer-
hmd. Zermatt, 9, $ (SMF); Walhs:

Frisch, 9, i (E. Schenkel, AMNH); Uri:
Susten Pass, 2232 m, 1875-1876, 9 (J. H.
Emerton, MCZ). France. Col de Portet
d'Aspet, juv. (E. Crosby, CU); Dept. Py-
renees Hautes: Lac d'Oo, 27 July 1932, 9
(E. Crosby, CU); Dept. Pyrenees Orien-
tales: Cerdagne many collections, 1976 (H.
Zibrowius, MCZ); Dept. Vancluse: Lu-
beron Mt., July 1976, 9 (H. Zibrowius,
MCZ); near Marseille, Aug. 1976, 9 (H.
Zibrowius, MCZ). Italy. Garda, 9, <5
(BMNH); Venice, 24 Aug. 1932, i (E.
Crosby, AMNH). Spain. San Lorenze de
El Escorial, 9 (CU). Yugoslavia. Ipek
|Pec|, 9 (SMF).

Aculepeira carbonaria (L. Koch)
Figures 198-207

Epeira carbonaria L. Koch, 1869, Z. Ferdinandeum,
Tirol, third ser., 14: 168. Female specimens from
Kiitthai, Finstertal, Tirol., believed lost.

Aranea carbonaria: — Roewer, 1942, Katalog der
Araneae, 1 ; 797.

Araneus carbonarius: — Bonnet, 1955, Bibliographia
Araneorum, 2(1): 451 (European records only).

This species is known only from Euro-
pean mountains.

Diagnosis. The coloration is dark (Fig.
203), resembling the American A. carhonari-
oides. Aculepeira carbonaria usually lacks
the paired light marks on the venter to the
side of the median white stripe ( Fig. 204 ) .
The scape of the epigynum is wider ( Figs.
198-201) than that of A. carhonarioides,
longer than that of A. ceropegia ( Fig. 200 )
and the median area of the base is white
(Figs. 199, 201); the palpal terminal apo-
physis tapers to the tip (Fig. 206), unlike
that of A. armida. The conductor is rela-
tively narrow with a deep distal depression
(Fig. 207); it differs from that of A. cero-
pegia but is somewhat similar to that of
A. armida.

Natural History. Aculepeira carbonaria,
as far as is known, is also limited to talus
slopes in the high mountains.

Specimens Examined. Austria. Tirol:
Stubai, 9, c^ (L. Koch, BMNH). Switzer-
land. Wallis: SaasTal, 9, S (E. Schenkel,

Metepeira, Kaiha, Aculepeira • Levi 235

Figure 217. Aculepeira armida (Audouin) (France). Epigynum. posterior view, cleared.

Figures 218-225. A. nosekl (Strand) (Turkey). 218-220. Epigynum: 218. Ventral. 219. Lateral. 220. Posterior.
221. Female abdomen, ventral. 222. Female, dorsal appendages removed. 223. Male left second patella and
tibia, dorsal. 224, 225. Male left palpus: 224. Mesal. 225. Ventral.

Figures 226-231. Aculepeira sp. (Samarkand). 226-229. Epigynum: 226. Ventral. 227. Lateral. 228. Posterior.
229. Posterior, cleared. 230. Female abdomen, ventral. 231. Female, dorsal, appendages removed.

Scale lines. 1.0 mm.

236 Bulletin Museum of Comparative Zoology, Vol. 148, No. 5

AMNII); Btm: Gommi Pass, 9 (J.H. Em-
erton, MCZ). France, "moiitagne," $ (E.
Simon, MCZ).

Aculepeira armida (Audouin),
new combination
Figures 208-217

Epeira armida Aiulouin, 1825 in Savigny. Explica-
tion somniaires dcs Planches d'Arachnides, in
Savigny, Description de I'Egyptc, p. 126, pi. 2,
fig. 8, 9 . Specimens from France, Italy and
vicinity of Acre I now northern Lsrael], lost. I
here re.strict the type locality to southern France.

Aranea armida: — Rocwer, 1942, Katalog der Ara-
neae, 1: 782.

Araneiis armidus: — Bonnet, 1955, Bibliograpliia
Araneoruni, 2(1): 439.

Note. This Mediterranean species has
not been found in America.

D/f/i^'/ios/.v. The species can be recognized
b\' the lobed dorsal abdominal band, en-
closing a dark mark and having a black
band on each side (Fig. 212). It differs
from the similar Neoscona adianto ( Walcke-
naer) by haxing a median ventral white
spot (Fig. 213); Neoscona has two or four
paired white marks on the venter. The
epigyniim differs from related species by
the relatively long scape (Fig. 210), by the
median lobe on each side of the base (Figs.
208, 209, 211), and by the thin elongate
lamellae bent at right angles behind the
base in posterior view (Fig. 211). The
palpus differs by having the tip of the ter-
minal apoph\'sis wider than the neck (Fig.
215), and by the shallow conductor, which
has distally a deeper area (Fig. 216).

Natural Histonj. The species has been
collected by sweeping prairies with isolated
tufts of high grass in Italy and from grazed
fields and bnish and cy|ir(>ss-pin(^ woods in
Yugoslaxia. Wiehle ( 1931 ) reports the spe-
cies from low brush or grasses with the
hub located, at most, at 50 cm above the
ground. The web has a white, silk-covered
hub and the retreat is flat, dish-shaped and
(;pen.

Collections Examined. Sicitzerland. Va-
lais: Valere near Sion, 9, ^ (NMB).
France. Luminy Col de Sugiton near Mar-

seille, 16 June 1976, 9 (H. Zibrowius,
MCZ). Spain. San Lorenz de Escorial, 9,
6 (CU); Sevdlla, 5 (NMB). Italy. Cala-
bria Prov. Aspromonte, 9 (E. Reimoser,
MCZ); Lazio Prov. Monteromano, 21 June
1965, S (P.M. BrignoU). Sardinia. 9, S
(SMF). Yufioslavia. Istria: 5 km north-
xx'cst of Vodrijan, 27 June 1962, 6 (H. & L.
Levi, MCZ); Dalmatia: Srebreno near Du-
brovnik, 13 June 1962, 9 (H. & L. Levi,
MCZ). Greece. 9 (BMNH). Tunisia. El
Kairovan, c^ .

Aculepeira noseki (Strand),
new combination
Figures 218-225

Araneus similis- Nosek, 1905, Ann. Kais. Konigl.
Naturhist. Hofmus. Wien, 20: 131. Numerous
female, male syntypes marked A. kariae from
Erdschias-Dagh [Ereiyas Dage Mountain] Tur-
key in very poor physical condition in Naturhis-
torisches Museum, Wien, examined. (Name pre-
occupied by Epeira similis Taczanowski, 1873
and Epeira simdis Bosenberg and Lenz, 1894).

Araneus noseki Strand, 1907, Zool. Jahrb. Abt. Sys-
tem., 24: 396. New name for A. sinnlis Nosek,
preoccupied.

Aranea kariae: — Roewer, 1942, Katalog der Ara-
neae, 1: 788. Roewer cites Strand, 1919, with-
out further citation.

Araneus karapagi Karol, 1966, Comm. Facult. Sci.
Univ. Ankara, 10(ser. C): 111, figs. 1-28, 9.
Female from Hakkari, Karada^, Turkey, in the
Museum National d'Histoire Naturelle, not ex-
amined. NEW SYNONYMY.

Diagnosis. The female differs from A.
armida in the more pronounced lobes of
the abdominal dorsal band, the coloration
is less contrasting (Fig. 222), and there
are straight lamellae that are posterodorsal
from the epigynum (Fig. 220); the male
differs by having a half-spear-shaped tip
of the terminal apophysis ( Fig. 224 ) .

Natural History. This species is also
found in talus at and above timberline ( K.
Thaler, personal communication ) .

Collections Examined. Turkey. Erzurum-
Paland()ken, 2400 m elev., 8 Aug. 1970, 9 ,
6 (V. Sbordone, PB); Altiparmek, 2400 m,
30 Aug. 1964, 9, S (KT); Lasistan, Cigu-
net Yayla, 2500 m, 20 Aug. 1968, 9 (H.

Metepeira, Kaira, Aculepeira • Levi 237

Gall KT); Verschambek, 2800 m, 11 Aug.
1965, 9 (H. Gall, KT).

REFERENCES CITED

Bonnet, P. 1957. Bihliographia Araneonim,

L'liiipiinierie Douladoure, Toulouse, 2(3):

1925-3026.
Chamberlin, R. V. AND W. IviE. 1942. A hun-
dred new species of American Spiders. Bull.

Univ. Utah, biol. sen, 7(1): 1-117.
CoMSTOCK, J. H. 1940. The Spider Book, rev.

edit. W. J. Gertsch, Doubleday, New York,

pp. 729.
Drensky, p. 1943. Die Spinnenfauna Bulgar-

iens. VI. Mitt, konigl. naturwiss. Inst. Sofia,

16: 219-2.54.
Karol, S. 1964. Sur une nouvelle espece du

genre Arancus ( Araneae, Argiopidae) origi-

naire d'Asie mineure. Bull. Mus. Nat. d'Hist.

Natur., Paris, 36 (ser. 2): 88-190.
. 1966. Une nou\elle espece du genre

Araneiis (Araneae, Argiopidae). Commun.

Facult. Sci. Univ. Ankara, 10 (ser. c): 111-

114.
Kaston, B. J. 1948. Spiders of Connecticut.

State Geol. Natiir. Hist. Surv., Hartford, Conn.

No. 70: 1-874.
Koch, C. L. 1834. in G. W. Herrich— Schiiffer,

Faunae Insectorum Germaniae, Regensburg,

Heft 126, p. 12, 13 (not seen, not available
to nie ) .

1839. in C. W. Hahn, Die Arachniden,

Niirnberg, 5: 1-158.
Le\i, H. W. 1977. The American orb-wcaNcr

genera Cyclosa, Mctuztjgia and Eusfala north

of Mexico (Araneae, Araneidae). Bull. Mus.

Comp. Zool., 148(3): 61-127.
LuBix, Y. D. 1975. Stabilimenta and barrier

webs in the orb-webs of Argiajje argentata

( Araneae, Araneidae ) on Daphne and Santa

Cruz Islands, Galapagos, J. Arachnol., 2:

119-226.
NosEK, A. 1905. Araneiden in A. Penther and

E. Zederbower, Ergebnisse einer naturwis-

senschaftlichen Reise zum Erdschias — Dagh

( Kleinasien). Ann. kais. konigl. naturhist.

Hofmus. Wien., 20: 114-154.
Roewer, C. F. 1942. Katalog der Araneae.

Verlag von Natura, Bremen, 1: 1-1040.
Strand, E. 1907. Spinnen des zoologischen In-

stituts in Tubingen. Zool. Jahrb., Abt. Syst.,

24: 391-468.
Thorell, T. 1870. Remarks on synonyms of

European spiders. Upsala. pp. 1-645.
Wiehle, H. 1931. Neue Beitrage zur Kenntnis

des Fanggewebes der Spinnen. Z. Moiphol.

Okol. Tiere, 22: 349-400.

238 Bulletin Museum of Comparative Zoolof^y, Vol. 148, No. 5

INDEX

N'alid nanu's are printed in italics. Pajfc num-
bers refer to main references; starred page num-
bers to illustrations.

aculeata, Epeira 228

Aciilcjjcira 222

Acidc})cira sp. 235*

alba, Caira 216

alba, F.peira 216

(ilba, Kaira 216, 217*

ahiventer, Kaira 218, 219*

alpina, Mctcpcira grandiosa 192*, 211*, 212, 213*

alpina, Metepeira 210

arizonica, Metepeira 199*, 200, 203*, 205*

armida, Acidepeira 233*, 235*, 236

armida, Aranea 236

armida, Epeira 236

armidus, Araneus 236

Caira 214

carhonaria, Acidepeira 233*, 234

earbonaria, Aranea 234

carbonaria, Epeira 228, 230, 234

carhonarioides, Aculepeira 224*, 229*, 230, 231*

earbonarioides, Epeira 230

earbonarius, Araneus 234

ceropegia, Aculepeira 229*, 231*, 234

eeropegia, Aranea 234

ceropegius, Araneus 234

eharitonovi, Aranea 230

comanchc, Metepeira 204, 207*

crassipes, Metepeira 190*, 202, 205*

crassipes, Metepeira 202

erueifera, Epeira 196

dakota, Metepeira 210

datomi, Metepeira 191*, 208, 209*

douglasi, Metepeira 198

ensenada, Metepeira 204

foxi, Metepeira 210, 211*

gogo.sa, Metepeira 200, 203*

, 212, 213*
214

212
203*

grandiosa alpina, Metepeira 192*, 211*

grandiosa grandiosa, Metepeira 213*,

gra)}diosa, Metepeira 210

grandiosa palustris, Metepeira 211*,

grinnelli, Metepeira 189*, 198, 201*,

hiteae, Kaira 219*, 220

hyperboreus, Araneus 230

inerma, Metepeira 208

jamaieensis, Metepeira 206

josepha, Metepeira 202

Kaira 214

karapagi, Araneus 236

kariae, Aranea 236

keyserlingi, Aranea 198

labyrintbea, Epeira 196, 210

labyrinthea grinnelli, Epeira 198

lahyrinfhea, Metepeira 188*, 196, 199*, 201*

Maepos 214

Metepeira 187

minima, Metepeira 206, 209*

nanella, Metepeira 210

noseki, Aculepeira 235*, 236

noseki, Araneus 236

obtusa, Kaira 218

packardi, Aculepeira 223*, 227*, 228, 229*, 231

paekardii, Epeira 228

palomara, Metepeira 210

palustris, Metepeira grandiosa 212

palustris, Metepeira 210

Pronaraebne 214

Pronarachne aries 219

sabino, Kaira 221*

septentrionalis, Araneus 228

similis, Araneus 236

spinosa, Caira 218

\'aehoni, Araneus 234

vegae, Araneus 230

Ventura, Metepeira 204, 207*

verae, Aculepeira 228

BuLietin OF THE

Museum of

Comparative

Zoology

Systematics and Zoogeography of Plagiola

[- Dysnomia = Epioblasma), an Almost Extinct

Genus of Freshwater Mussels (Bivalvia:

Unionidae) from Middle North America.

RICHARD I. JOHNSON

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148 NUMBER 6
4 APRIL 1978

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chusetts, 02138, U.S.A.

SYSTEMATICS AND ZOOGEOGRAPHY OF PLAGIOLA
(= DYSNOMIA = EPIOBLASMA), AN ALMOST EXTINCT
GENUS OF FRESHWATER MUSSELS (BIVALVIA:
UNIONIDAE) FROM MIDDLE NORTH AMERICA.

RICHARD I. JOHNSON'

CONTENTS

Abstract 239

Introduction 240

Background 240

Rele\ant Faunal Studies 241

Zoogeograph\- 242

General Consideratioxs 242

Distribution of Plagiola Below the

Area of NLaximum Glaciatiox 243

Post-glacial Distributiox of Plagiola __ 244

Obser\ations and Conclusions 244

Acknowledgments - 245

Systematic Section 246

AhJ}icviations 246

SynoniiDiy 246

Desc: iptioiis 246

Anatomy and Breeding Season 246

Habitat 246

Remarks 246

Range 246

Abundance 246

Specimens examined 246

Distribution 246

Figures i 247

Key to the subgenera of Plagiola 247

Genus Plagiola Rafinesque 247

Subgenus Truncillopsis Ortmann and Walker 248

Plagiola (T. ) triquetra (Rafinesque) 248

Subgenus Plagiola S. S. Rafinesque 252

Plagiola (P.) internipta (Rafinesque) 252

Plagiola (P.) penita (Conrad) 254

Plagiola (P.) arcaeformia (Lea) 257

Plagiola (P.) lenior (Lea) 259

Subgenus Torulosa Friersor 260

Plagiola (T.) tondosa (Rafinesque) 261

Plagiola (T. ) sampsoni (Lea) 265

Plagiola ( r. ) propinqua (Lea) 266

Plagiola ( T. ) biemarginata ( Lea ) 268

Plagiola ( T. ) capsaeformis ( Lea ) 269

Plagiola (T.) florcntina (Lea) 271

Plagiola (T.) turgidula (Lea) 274

Subgenus Pilea Simpson 276

Plagiola ( P. ) personata ( Say ) 276

Plagiola (P.) obliquata (Rafinesque) 278

Plagiola (P.) haysiana (Lea) 280

Subgenus Epioblasma Rafinesciue 282

Plagiola (E.) flexuosa (Rafinesque) 283

Plagiola (E.) stewardsoni (Lea) 285

Literature Cited 287

Index to Rele\'ant Taxa 291

^ Museum of Comparative Zoology, Harvard
University 02138.

Abstract. Plagiola ( — Dysnomia = Epioblas-
ma), a genus of Unionidae (Mollusca: Bivalvia)
is unique among all freshwater mussels in the ex-
tent of sexual dimorphism found among its mem-
bers. The 17 species recognized here are assigned
to 5 subgenera. All but one species, found in the
Mobile-Alabama-Coosa river system in Alabama,
occur in the Tennessee Ri\er system, and that spe-
cies is clearly deri\ed from one in the latter sys-
tem. Fourteen of the species are also found in the
Cumberland Ri\er s>stem. Three of tliese fourteen
are also found in the \\"hite Ri\er system on the
Ozark Plateau in Missouri and Arkansas. These
three species, as well as a number of other simi-
larly distributed unionids, afford evidence of a

Bull. Mas. Comp. Zool., 148(6): 239-320, April. 197S 239

240 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

relict fauna that may have persisted since the
Cretaceous. The post-glacial distribution of several
of the species that ha\e found their way beyond
the Tennessee and Cumberland ri\er systems sug-
gests that all the species are of Cumberlandian
origin.

INTRODUCTION

Among the numerous genera of Uni-
onidae, Plagiola is one of the more interest-
ing because of the extent and variety of
sexual dimorphism among the 17 species
belonging to 5 subgenera. Many of the
species vary but slightly, though several
have numerous ecophenotypic varients.
Some forms, formerly recognized as valid
species or subspecies, could not be properly
dealt with before the biological species
concept was defined. This paper attempts
to clarify the synonymy of the species of
PhiiS,iola and to reconstruct what was, until
a short time ago, their distribution. To do
this, most specimens in the principal mu-
seums were examined. These records are
supplemented with some only available in
the literature. Almost a dozen of the spe-
cies are now considered extinct or endan-
gered from pollution, or because their habi-
tats have been, or are being, destroyed by
impoundments.

BACKGROUND

The members of Plagiola were first mono-
graphed and separated from Unio by
Simpson (1900a: 524), who placed them
in the genus Tmncilla Rafinesque 1820.
His designation of Tmncilla triqueter Rafin-
esque as the type was invalid, since Tmn-
cilla tmncata Rafinesque had previously
been designated as the type by Hernnann-
sen (1849, 2: 627). Therefore, Ortmann
and Walker (1922: 65) designated Dys-
noniia Agassiz 1852 as the next available
generic taxon, whose type species was Unio
foliatus Hildreth (= [Ohliquaria] flexiiosa
Rafinesque) by subsequent designation
(Simpson 1900a: 521). They made this
designation since they regarded the type
species of both Plagiola Rafinesque 1820
(Ohliquaria (Plagiola) intermpta Rafin-

esque) and Epiohlasma Rafinesque 1831
(E. hiloha Rafinesque) as unidentifi-
able, though Frierson (1914) had previ-
ously asserted that hiloha was the female
oi foliatus Hildreth. Thiele (1934), Clench
(1959), Morrison (1969) and Stansbery
(1973), used Epiohlasnui over Dijsnomia.
Stansbery (pers. com.) assured me that he
can recognize hiloha. To settle this matter,
a neotype was chosen for it here, see p. 283.
However, Johnson and Baker's (1973:159)
selection of a lectotype for Ohliquaria (Pla-
giola) interrupta Rafinesque, which Morri-
son (1969) had previously asserted as iden-
tifiable, makes Plagiola the earliest
available generic taxon, with Epiohlasma
(—Dijsnomia) as a subgenus.

These nomenclatorial changes are un-
fortunate, but we now have Rules of Inter-
national Nomenclature (1964), not avail-
able to Ortmann and Walker, which
emphasize the identification of the type. As
Stansbery (pers. com.) has said, "Ortmann
and Walker (1922) did a fine job for their
time in consideration of their basic premise
that the validity of a name depends upon
its identifiability from the original descrip-
tion. I consider the identification of the
holotype (if extant) to be the court of last
appeal in such matters. If we did not, most
of Lamarck's [unionid] names now in use
would become nomina duhia."

The most recent revision of the Rules
[1974. Bull. Zool. Nomencl. 31 (2): 80]
under Article 2-3 states, "A zoologist who
considers that the application of the Law
of Priority would in his judgment disturb
stability or universality or cause confusion
is to maintain the existing usage and must
refer the case to the Commission for a de-
cision under the plenary powers [Art. 79]."

It is hoped that in this paper, the identity
of the specific taxa of Plagiola are settled.
It is unfortunate that some of these identifi-
cations also result in changes on the ge-
neric and subgeneric level. However, Mor-
rison (1969) declared Plagiola available
over Dijsnomia, and Stansbery has used
Epiohlasma over Dijsnomia numerous times

Plagiola from Middle North America • Johnson 241

in the literature. Since this author is not
sure what is existing usage in Plagiola, this
paper is written on the assumption that the
Law of Priority has not been totally ab-
rogated.

Simpson (1900a: 516-524) recognized
four subgenera within this genus. Ortmann
and Walker (1922: 65) created monotypic
Truncillopsis, based on the former's studies
of the very primitive anatomy of triquetra.
Frierson (1927: 93-96) added three addi-
tional subgenera. These were based en-
tirely on conchological characters, merely
by selecting a type species for each, with-
out explanation. In spite of this method,
two of his subgenera are recognized here,
viz. Plagiola Rafinesque s. s. [=Penita
Frierson] and Torulosa Frierson {=Capsae-
formis Frierson].

Walker ( 1910 ) constructed an excellent
key to the species, as he conceived them.
Simpson (1914) augmented his previous
arrangement ( 1900a ) with species descrip-
tions. Haas (1969: 477^90), in Das Tier-
reich, wrote on Dijsnomia [—Plagiola]. His
subgeneric and specific concepts are those
of Frierson, and his descriptions appear to
be essentially German translations of Simp-
son (1914: 2-32). Haas's work is a com-
pilation of the literature on this genus to
1927. Burch (1973, 1975) included the spe-
cies as recognized by Simpson in a general
key to the North American Unionacean
clams.

RELEVANT FAUNAL STUDIES

The unionid fauna of the upper Tennes-
see River system was commented on by
Coker (1912), studied extensively by Ort-
mann (1918), and reexamined by Stans-
bery (1973) and Stansbery and Clench
( 1975 ) . The lower Tennessee, below Wal-
den Gorge, to the Muscle Shoals in Lauder-
dale and Colbert counties, Alabama, was
extensively studied by Ortmann (1925)
and augmented with notes by van der
Schalie (1939b). Morrison (1942) com-
pared the fauna found in the Indian
mounds near the Muscle Shoals with the

present fauna in the river. Isom ( 1968 )
and Isom and Yokley (1968) enumerated
the unios of Indian Creek and Bear Creek
tributaries of the Tennessee in Alabama.
Duck River, a tributary of the lower Ten-
nessee, was carefully studied by Ortmann
(1924a), restudied by Isom and Yokley
(1968) and re-restudied by van der Schalie
(1973).

The Cumberland River unionids were
studied by Wilson and Clark (1914).
Those of the upper part of the river were
examined again, between 1947^9, by Neel
and Allen ( 1964 ) before the completion of
the Wolf Creek Dam.

Studies have been made of the unionid
faunas of many of the rivers flowing into
the southern side of the Ohio River beyond
the Cumberland. These are presented in
a west to east arrangement.

The unios of the Tradewater River were
listed by Clench and van der Schalie
(1944); those of the Green were exten-
sively studied by Ortmann (1926). The
Green River species were again listed by
Clench and van der Schalie (1944), who
also included the Salt River unionids in
their paper. The Salt River unionids were
listed again by Rosewater (1959). The
unionids of the Kentucky River were stud-
ied by Danglade (1922), and Ortmann
(1913: 290) discussed the species found in
the upper Ohio drainage.

Relevant studies of the Ohio River union-
ids and of those rivers flowing into it on
the northern side are presented in a west
to east arrangement. The unionid fauna of
Kansas was studied by Scammon (1906) and
again by Munay and Leonard (1962). Ut-
terback (1915-16) studied that of Mis-
souri, and recently Buchanan (ca. 1976)
studied that of the Meramec River basin in
the same state. Baker ( 1928 ) wrote on the
unionids of Wisconsin. Parmalee ( 1967 ) , in
a popular paper on the Unionidae of Illi-
nois, included data on Plagiola, as did Star-
rett (1971) in his work on the Unionacea
of the Illinois River. Stein ( 1881 ) enumer-
ated the mollusca of Indiana; these were

242 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

monographed by Call (1900). Call's work
was supplemented bv Blatchley and Dan-
iels (1903) and Daniels (1915). The In-
diana mollusca were subsequently revised
by Goodrich and van der SchaHe (1944).
Meyer (1974) studied the unionid fauna of
the ^^'abash and ^^■ hite rivers in Indiana,
and Clark ( 1976) examined the unionids of
the lower Wabash River. Baker (1922)
had previously studied the molluscan fauna
of the Big Vermilion River, a large tribu-
tary of the Wabash. Goodrich (1932)
wrote a handbook on the mollusca of Michi-
gan and included data on PlogioJa. La
Rocque (1967) compiled a work on the
Unionidae of Ohio.

Wilson and Clark (1912b) reported on
the extensive collection of naiades they
made in the Maumee River drainage, which
is tributary to Lake Erie. Clark ( 1977 )
wrote on the naiades of the St. Joseph River
of the Maumee. Ortmann (1924b) in a pa-
per on the distributional features of the
naiades in the tributaries of Lake Erie, dis-
cussed the post-glacial dispersal of the
species, as did Goodrich and \'an der
Schalie ( 1932 ) , who studied the unionids
of the Great Lakes. The post-glacial dis-
persal of unionids to Lake Erie was again
reviewed by Stansbery (1961).

\'an der Schalie made further contribu-
tions to the understanding of post-glacial
dispersion in his papers on the unionid
faunas of the Muskegon, Grand (both
1941a), St. Joseph (1936) and Huron
(1938a) river drainages in Michigan. In
a paper discussing the value of mussel dis-
tribution in tracing stream confluence, van
der Schalie (1945) summarized the data
about the post-glacial dispersal of the
Unionidae.

The unionid fauna of several drainages
of the Nh^bile-Alabama-Coosa river system
have been studied. The unionids of the
Cahaba and Tombigbee rivers were ex-
amined by van der Schalie (1938b, 1939a),
and those of the Coosa River by Hurd
(1974).

ZOOGEOGRAPHY
General CoNsroERAxioxs

It should be mentioned to the reader
unfamiliar with the means of dispersal of
the Unionidae, that their mobility is pas-
sive— dependent on fishes to which the lar-
val forms, or glochidea, attach themselves
for a period of time. Stream capture, and
subsequent rupturing of confluences, have
therefore played a significant role in de-
termining the geographic distribution of
the Unionidae.

The Tennessee and Cumberland river
systems are among the world's most an-
cient. The Tennessee, containing at least
86 species of Unionacea, has the largest
assemblage of unionid species found any-
where, followed by the Cumberland River,
which has a unionid fauna of at least 78
species.

Ortmann (1924a: 40) recognized among
the Unionidae two distinct faunal elements
in these two river systems — those belong-
ing to the Interior Basin and found in the
Ohio River drainage, and those not found
outside the Tennessee and Cumberland
river systems. (There are a few exceptions
that also occur on the Ozark Plateau ) . This
latter group of species is restricted largely
to the Cumberland Plateau and to the Great
Allegheny Valley. This area represents the
Cumberlandian Region, defined by Ort-
mann (1924a: 40) as including only the
drainages of the Tennessee River system
from the headwaters to the vicinity of Mus-
cle Shoals, in Colbert and Lauderdale coun-
ties, Alabama; and the Cumberland River
system from the headwaters to the vicinity
of Clarksville, Montgomery County, Ten-
nessee (Ortmann, 1925: 366). Ortmann
(1924a: 40) also discussed the unionid
fauna of the Duck River drainage, which is
at present a tributary of the Tennessee
River system. The upper portion of the
Duck River has a fauna that is 38 per cent
Cumberlandian, and Ortmann suggested
that this was the original fauna and that
there had been stream confluence with the

Plagiola from Middle North America • Johnson 243

Duck and botli the Tennessee and Cumber- Another species, Ciimherlandia mono-
land river systems long ago. donta (Say), of the family Margaritiferidae,
The distribution of the old Interior Basin is found in both the Tennessee and Cum-
fauna in the Tennessee and Cumberland berland river systems and north of the
river systems indicates that most of it was Ozark Crest in the Osage and Gasconade
present in these rivers, along with the even rivers in Missouri. Like triquetra, it ap-
more ancient Cumberlandian fauna, long pears to have had refugia here from glacial
before maximum Pleistocene glaciation destruction. The members of CAjprogenia
( which occurred early in the epoch and ex- and a number of other unionid species
tended southward roughly to the present have the same restricted distribution as
Missouri and Ohio rivers). Apparently many florentina and turgidula, but they are be-
members of the former fauna, and some of yond the scope of this paper,
the latter, repopulated at least the present Seven species of Plagiola — interrupta
Ohio and upper St. Lawrence river systems (Plate 7, fig. A), capsaeformis (Plate 7, fig.
from these sources. Some of the species B), /enior (Plate 7, fig. C), /jof/.siflnfl (Plate
may also have had refugia elsewhere, as in <S, fig. A ) , arcaeformis ( Plate 8, fig. B ) ,
the Allegheny and Monongahela rivers in stewardsoni (Plate 8, fig. C) and hieniar-
Pennsylvania (Ortmann, 1912b). ginata (Plate 9, fig. A) — are found in both
Originally Ortmann (1924a: 40) regarded the Tennessee and Cumberland river sys-
as Cumberlandian only those unionid spe- tems exclusively.

cies currently confined to the areas of the Plagiola penita of the Mobile-Alabama-
Tennessee and Cumberland river systems, Coosa river system is derived from inter-
as defined above. Later ( 1925: 370) he sug- rupta of the Tennessee River system,
gested there were Cumberlandian species Hayes and Campbell (1894) suggested that
that descended the Tennessee and Cumber- the upper Tennessee River fomierly flowed
land river systems and invaded the Ohio through WMen Gorge into the Gulf of
River drainage and the Interior Basin. ^^xico by way of the present Mobile-Ala-

bama-Coosa river system, and that it was

Distribution of Plagiola below diverted to its present course through the

THE AREA MAXIMUM Glaciation Cumberland Plateau in the late Tertiary.

Their conclusions, based entirely on physio-
All of the species of Pkgiola, with the ex- graphic evidence such as the character of
ception of penita from the Mobile-Ala- the Tennessee-Coosa divide, were substanti-
bama-Coosa river system in Alabama, ated by Simpson (1900b) on the basis of
occur in the Tennessee River system, and similarities in the unionid faunas of the
only two of these, torulosa ( Plate 3 ) and present river systems.

sampsoni (Plate 6, fig. B) are missing in Johnson (1905) effectively defeated all
the Cumberland River system. support for the river capture theory of
Three species, triquetra (Plate 1), tur- Hayes and Campbell. The upper Tennes-
gidtda and florentina (both Plate 2), are see appears to have held the same course
found not only in the Tennessee and Cum- since the close of the Cretaceous. However,
berland river systems, but also in the White "where the smaller tributaries of the Coosa
River system in Missouri and Arkansas, and Tennessee rivers have common divides
south of the Ozark Crest, where they may in the carbonate rocks of the Appalachian
have persisted since before the Cretaceous Valley, it is highly probable that many cap-
uplift. One of these species, triquetra, is tures have occurred and effected the faunal
also found north of the Ozark Crest in the transfers which have been formerly at-
Meramec River, Missouri, and in tributaries tributed to the Walden Gorge capture."
of the present Missouri River in Kansas. (Hurd, 1974: 137).

244

Bulletin Museum of Comparative Zoologij, Vol. 148, No. 6

Post-glacial Disthibution of Plagiola

Maximum Pleistocene glaciation occurred
earl\- in the epoch, and extended southward
to roughly the present Missouri and Ohio
rivers, \iuch of the unionid fauna found
on the Cumberland Plateau is the same as
that found on the Ozark Plateau, and it is
assumed that the latter area was as an im-
portant source for the species that repopu-
lated the Mississippi drainage (above the
Missouri ) , as was the former in the repopu-
lation of the Ohio drainage. Ortmann
(1913: 351) obsei-ved that the unionid
fauna of the Ohio River is more numerous
downstream. There are some 60 species in
the vicinity of Cincinnati, Hamilton County,
Ohio, decreasing to 47 species in Pennsyl-
vania. Ortmann suggested that this fauna
migrated upstream in glacial and post-
glacial time when the present Ohio River
was formed. The unionid fauna of the
larger tributaries of the Ohio River drain-
age in Kentucky, beyond the limit of glaci-
ation, is Ohioan. From west to east, the
tributaries are: the Tradewater with 17
species, the Green with 50 species, the Salt
with 22 species {all Clench and van der
Schalie, 1944), the Kentucky with 40 spe-
cies (Danglade, 1922), the Licking with 14
species and the Big Sandy with 12 {both
Ortmann, 1913).

Plagiola triquetra is the only member of
the genus found in the Mississippi River
drainage. While it may have spread there
from the Ohio River, it is just as likely
that it spread from a refugium in the Mera-
mec River system, Missouri. In any event,
the present distribution of triquetra in Wis-
consin clearly illustrates van der Schalie's
(1945: 336) suggestion that a connection
existed in post-glacial time between the Fox
and Wisconsin'rivers at Portage, Columbia
County, Wisconsin (Plate 1, A).

The presence of triquetra in the Illinois
River, Illinois, and in the Muskegon, Grand
and St. Joseph rivers, on the eastern side of
Lake Michigan in the St. Lawrence drain-
age, supports van der Schalie's (1945: 356)
suggestion that before the formation of

Lake Michigan the latter rivers were tribu-
taries of the Des Plaines River, which by
way of the Chicago outlet, drained into the
Mississippi River (Plate 1, B). Although
torulosa is not currently found in the Mis-
sissippi River drainage, the species prob-
ably reached the Grand River by the same
route as that taken by triquetra (Plate 3).
If it ever occurred in the Cumberland River
system, it now appears to be missing.

It is assumed that triquetra spread into
the Ohio River system from the Tennessee
and Cumberland river systems, though it
might have also had refugia in the Alle-
gheny and Monongahela river drainages in
western Pennsylvania (Ortmann, 1912b),
Fig. 1 ) . The distribution of triquetra as well
as torulosa and ohliquata in the St. Law-
rence River system indicates a former con-
nection between the Wabash and Maumee
rivers in the vicinity of Fort Wayne, Allen
County, Indiana (Plate 1, C, Plate 3, Plate
4). Like triquetra, torulosa might have had
refugia in western Pennsylvania.

The present distribution of triquetra, to-
rulosa and ohliquata in the rivers flowing
into Lake Erie also indicates the correct-
ness of Ortmann's (1924b) and van der
Schalie's (1945: 362) view that during the
Trent Oudet stage of the Great Lakes, when
Lake Erie was partially dry, many of the
present rivers flowing into western Lake
Erie were part of the Greater Maumee
River system.

Three species, flexuosa (Plate 5), per-
sonata (Plate 6, fig. A), and propinqua
(Plate 6, fig. B) have spread from the
Tennessee and Cumberland river systems
into the Wabash River drainage and the
Ohio River. An additional species, samp-
soni (Plate 6, fig. B), missing from the
Cumbcriand River, has a restricted post-
<dacial distribution similar to flexuosa and
personata.

OBSERVATIONS AND CONCLUSIONS

1. There are 17 species of Plagiola; all
but one occur in the Tennessee River sys-
tem. P. penita of the Mobile-Alabama-

Plagiola from Middle North America • Johnson 245

Coosa river system is clearly derived from
P. inferrupta and provides evidence of a
former confluence between the two river
systems in the past.

2. The Cumberland River system has 14
of the 16 species found in the Tennessee
River system, lacking only P. torulosa
and sampsoni. Obviously, there has been
stream confluence between these ancient
river systems.

3. There are six species of Tla^iola in the
upper Duck River drainage of the Ten-
nessee River system; only one, P. triqtietra,
is found outside the Tennessee or Cumber-
land river systems. Ortmann (1924a: 46),
on the basis of this and much more data,
suggested that Duck River was originally
more directly connected with the Tennes-
see and Cumberland rivers.

4. Two species of Plagiola, tttrgiduJa and
florentina, occur only in the upper Ten-
nessee and Cumberland river systems and
in the upper W^hite River system south of
the Ozark Crest in Missouri. The presence
of these species in the latter system as well
as P. triquetra and several other unionid
species (not discussed in this paper),
strongly suggests that a number of species
have persisted since the Cretaceous uplift.

5. Seven species of Plagiola, found in the
Tennessee River system, have spread into
the fonnerly glaciated area. Two of these,
P. triquetra and torulosa, may have had
Pleistocene refugia in the Allegheny and
Monongahela river drainages in the moun-
tainous region of western Pennsylvania.
Plagiola torulosa and sampsoni do not
occur in the Cumberland River system.
Thus only five species of Plagiola: triquetra,
propinqua, personata, obliquata and flexu-
osa, may ha\'e spread into the once glaci-
ated area from the Cumberland River sys-
tem.

6. The present distribution of P. tri-
quetra in Wisconsin suggests that a con-
nection once existed in post-glacial time
between the Fox and Wisconsin rivers at
Portage, Columbia County, Wisconsin.

7. The present distribution of P. triquetra

— in the IlHnois River, Illinois, and the
Muskegon, Grand ( as well as torulosa in the
latter) and St. Joseph rivers on the eastern
side of Lake Michigan in Michigan — indi-
cates that before the formation of Lake
Michigan, the latter streams were tribu-
taries of the Des Plaines River, which
drained into the Mississippi River by way
of the Chicago outlet.

8. The present distribution of P. tri-
quetra, torulosa and obliquata in the St.
Lawrence River system indicates that a
connection fonnerly existed in post-glacial
time between the Wabash and Maumee
rivers in the vicinity of Fort Wayne, Allen
County, Indiana.

9. The present distribution of P. tri-
quetra, torulosa and obliquata in the rivers
flowing into western and southern Lake
Erie indicates that during the Trent Outlet
stage of the Great Lakes, when the bed of
Lake Erie was partially dry, these rivers
were part of the Greater Maumee River
system.

10. The most primitive species of Pla-
giola, triquetra, appears to be the most
abundant as well as the most widely dis-
tributed species in the genus. Interestingly,
the shells exhibit little moiphological vari-
ation.

11. Following Plagiola triquetra, the
most widely distributed species in the genus
are torulosa and obliquata, in that order.
The shells of torulosa exhibit considerable
ecophenotypic variation depending on their
environment, while those of obliquata show
almost no such variation.

12. Three species of Plagiola, arcaefor-
mis, stewarcliana and biemarginata, appear
to be the least abundant as well as the least
widely distributed species in the genus.
The shells exhibit little morphological vari-
ation. All three species are now considered
extinct.

ACKNOWLEDGEMENTS

Special thanks are extended to Dr. Joseph
Rosewater, National Museum of Natural
History, who, in addition to giving me ac-

246 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

cess to the collection, had photographs
made of all of the relevant types, and to Dr.
Arthur H. Clarke for the loan of material.
Dr. George M. Davis, Academy of Natural
Sciences of Philadelphia, made that collec-
tion available and generously loaned types
and specimens. Dr. Juan J. Parodiz, Car-
negie Museum, Pittsburgh, kindly made
that collection available for study. Drs.
Henry van der Schalie, John B. Burch and
Charlotte Patterson, and Mr. R. W. Hanley,
Museum of Zook)gy, University of Michi-
gan, made the collection in their charge
available. Dr. Alan G. Solem, Field Mu-
seum, Chicago, graciously loaned specimens
and answered questions. Thanks are also
extended to Dr. David H. Stansbery, Ohio
State Museum, for several data on Plogiola,
a genus that is also of interest to him; to
my colleague, Dr. Kenneth J. Boss; and to
the re\'iewers, for reading the manuscript
and making significant suggestions.

SYSTEMATIC SECTION

Ahhrev)iatio)\s. The following abbrevia-
tions have been used in the text and on the
plate captions:

ANSP Academy of Natural Sciences of
Philadelphia, Pennsylvania

BMNH British Museum (Natural History),
London, England

CM Carnegie Museum, Pittsburgh, Penn-

sylvania

MCZ Museum of Comparative Zoology,

Cambridge, Massachusetts

MZUM Museum of Zoology, University of
Michigan, Ann Arbor, Michigan

OSM Ohio State Museum, Columbus, Ohio

USNM National Museum of Natural History,
Washington, D. C.

Synonymy. For ease of reference, full
citations are included for each taxon, in-
cluding the type locality and the location of
the type when known. References to plates
and figures are not included under Lea's
Obs. Unio since they are always the same
as in the preceding entry. In some in-

stances, lecto types are selected. Elsewhere
in the text, references are abbreviated and
require the use of the bibliography. Ex-
cept for the original references, only rele-
vant citations since 1914 are included here
since the earlier ones are available in Simp-
son (1914).

Descriptions. The measurements are
only intended to convey the general size of
specimens from a given station, and to in-
dicate sexual differences.

Anatomy and Breeding Season. The
available data are cited.

Habitat. Included when known.

Remarks. These are designed to elucidate
the differences between the sexes and the
species, and include comments on distri-
bution and taxonomy.

Range. The distribution is summarized.

Abundance. The former abundance of
the species is based on the number and size
of the lots found in the several collections
studied. Their present abundance is based
on the opinions of Stansbery (1970, 1971,
1976).

Specimens Examined. Most of the rec-
ords are based on the specimens in the col-
lections mentioned above. These collections
contain almost all specimens available for
study, and with the exception of the small
collection in the Ohio State Museum, all
have been personally examined. The rec-
ords are followed by the initials of the
institution in which they were observed.
In most instances when records were du-
plicated only references to the specimens in
the Museum of Comparative Zoology were
included, though in certain critical cases
several references are mentioned. Speci-
mens not seen are credited to the responsi-
ble individual or published reference and to
the associated institution, if it is known.
The published references may be found
either under Synonymy or under Literature
Cited, or sometimes under both headings.

Distribution. The various river systems
are listed from west to east. East of the
Mississippi River this arrangement approxi-

Plagiola from Middle North America • Johnson 247

mates the direction of post-glacial dispersal.
The records from each river system are ar-
ranged from the headwaters to the mouth,
with the exception of the Ohio River drain-
age, which is oppositely arranged to indi-
cate the post-glacial distribution of most
species from the Tennessee and Cumber-
land river systems.

The cumbersome term, Mobile-Alabama-
Coosa river system, is used because it re-
flects the main channel of the river, which
was given a different name at each impor-
tant confluence.

Figures. When available, holotypes, al-
lotypes and lectotypes are generally used
to illustrate each species. Where required,
several illustrations are included to show
ecophenotypic variation. Not included are
the pencil sketches of the new species de-
scribed by the Sicilian nobleman, Marchese
A. De Gregorio ( 1914), who, with abandon,
began redescribing the unionids of North
America. Some of the data on the plate
captions, such as the measurements, are not
repeated elsewhere.

The distributional maps are based on
Rand McNally and Company's Commercial
Atlas of America 1912 Edition, plate 21,
which indicates United States Inland Wa-
terways.

Key to the Subgenera of Plagiola

Because of the pronounced sexual dimorphism
occurring in Plagiola, this key to the subgenera con-
sists of one section for each sex. The keys to the
species in the several subgenera are constructed
without special regard for sexual differences. Any-
one unfamiliar with this genus should refer to
Burch's "Key" (1973 or 1975).

Males

1. Shell with a distinct radial furrow in front of

the posterior ridge 2

Shell without a distinct radial furrow in
front of the posterior ridge 4

2. Radial furrow narrow and shallow Pilea

Radial furrow wide and of varying depth 3

3. Shell subrhomboid or subquadrate

Epioblasma

Shell oval, obovate, elliptical, subquadrate
or trapezoid ToitiJosa

4. Shell long based triangular ._ Truncillopsis

Shell subquadrate, subrhomboid, or elliptical
Plagiola

Females

1. Marsupial expansion occupying part of the

posterior or medial region 2

Marsupial expansion occupying the entire
postbasal region _____ Torulosa

2. Marsupial expansion formed by a swelling of

the posterior ridge _ 3

Marsupial expansion formed by a swelling in
front of the posterior ridge and more or less
separated from it 4

3. Shell distinctly long based triangular, mar-
supial swelling not extending below the base
line, with conspicuous green rays and mot-
tling TrunciUo})sis

Shell subquadrate or subrhomboid, marsupial
swelling usually extending below the base
line, usually with fine green rays that are
often broken by growth rests Plagiola

4. Marsupial area located medially and extend-
ing below the base line as a distinct lobe

Epioblasma

Marsupial area in front of the posterior ridge,
often separated from it by a narrow sulcus,
not extending below the base line Pilca

Superfamily UNIONACEA Thiele 1935

Family UNIONIDAE (Fleming 1828) Ort-

mann 1911

Subfamily LAMPSILINAE (Ihering 1901)

Ortmann 1910

Genus Plagiola Rafinesque

Plagiola Rafinesque 1819, Jour. Phys. Chim. Hist.
Nat. (Paris) 88: 426. Species listed: verrucosa,
fasciolaris, leptodon, depressa, flava, obliquatas
[sic], all Rafinesque, all iiomina nuda; 1820, Ann.
Gen. des Sci. Physiques, Bruxelles 5: 302.
Species listed: Obliquaria dccorlicata, O. inter-
nipta, O. depressa, O. lineolata, all Rafinescjue.
Type species, Vnio interrupttis Rafinesque 1820,
subsequent designation, Herrmannsen, 1847,
Indicis Generum Malacozoorum 2: 279.

Penita Frierson 1927, Check list N American na-
iades, pp. 11, 93. Type species, Unio penitus
Conrad, original designation, teste Errata et
Corrigenda.

Description. The most interesting char-
acter of Plagiola is the remarkable differ-
ence between the shells of the male and fe-
male. The shells are essentially alike until
about one-third to one-half grown, when
the development of marsupial swelling be-
gins in the female. This marsupial swelling,

248 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

in the post-ventral region, is thinner than
the rest of the shell and is often of a some-
what different texture; it may take the form
of a widely rounded wing, or be marked
off from the rest of the shell by sulci. The
swelling is often radialh' sculptured and
toothed at the edge, with remains of the
teeth visible along the growth lines. In
many species the male has a radial de-
pression in front of the posterior ridge. The
shape of the shell is variable, but is gener-
all\- suboN'ate or subtriangular and is some-
what inflated. The shell is either not sculp-
tured or only covered with low tubercles.
The umbonal sculpture is delicate and
faintly doubly looped. The periostracum is
yellowish or greenish, generally with some
green rays.

Anatomy. Discussed by Ortmann
(1912a: 354), Simpson (1914: 2) and Utter-
back (1916: 452 [189]).

The selection of a lectotype for Ohli-
quciria {Plagiola) interruvta Rafinesciue
1820 by Johnson and Baker (1973: 159)
and its acceptance in this paper over Unio
hrevidens Lea 1831 requires that the spe-
cies, formerly under the genus Dysnomia
or Epiohlasma, be placed in Plag^iolo. This
use of Pla^iola was indicated by Morrison
(1969:24).

Ortmann and Walker (1922: 51) re-
garded O. (P.) intenupta Rafinesque as
unidentifiable and designated O. (P.) Une-
olata Rafinescjue = securis Lea as the type
species of P]a<i,io]a. This apparently made
Dysnomia Agassiz 1852 available. See un-
der remarks to subgenus Epiohlasma.

Baker (1964: 140) pointed out that
whether or not interrupta was identifiable,
Ortmann and W^alker's subsequent designa-
tion of Uneolata as type species was in-
valid, and that Uneolata belonged in the
genus EUipsaria Rafinesque, the synonymy
of which follows:

Ellipsaria Rafinesque 1820, Ann. Gen. des Sci.
PliNsiques, Braxelles 5: 303. Type species, Ohli-
qiiaria ellipsaria Rafinesque, by tautonymy. Ort-
mann and Walker (1922: 52) aj;(rced that tliis
ta.xon is a synonym of Uneolata. Their conclu-
sion was confirmed by Johnson and Baker

(1973: 154, pi. 5, fig. 1), who selected a speci-
men identified by Rafinesque as neotype.

Crcnodonta Schliiter 1838, Kurzgefasstes systema-
tisches Verzeichness meiner Conchyliensamm-
lung .... (Halle), p. 33. Species listed:
plicata Say, tuherculata Rafinesque, securis De-
shayes [=Lea], trigona Lea. Type species,
Crenodonta securia (Deshayes) [=Lea], subse-
quent designation, Herrmannsen, 1852, Indicis
Generum Malacozoorum, Supplementa et Cor-
rigenda, p. 38.

Plagiolopsis Thiele, 1934, Handb. syst. Weich-
tierk. 3: 834. Type species, P. securis (Lea),
monotypic.

Thiele concluded that Plagiola was
equivalent to Lanipsilis Rafinesque and, un-
aware of either Ellipsaria Rafinesque or
Crenodonta Schliiter, created an unneces-
sary taxon.

Ellipsaria is monotypic, and is repre-
sented by E. Uneolata Rafinesque 1820.

Subgenus Truncittopsis Ortmann and
Walker

Truncillopsis Ortmann and Walker 1922, Occ. Pa-
pers, Mus. Zool., Univ. Mich. no. 112, p. 65.
Type species, Truncilla triqiieter Rafinesque,
original designation.

Description. Shell long based triangular,
covered with conspicuous green rays and
mottling, greatly inflated, sharply tnmcated
posteriorly. Disk smooth without any radial
furrow. Female with a slight marsupial
swelling at the posterior ridge ending at
the extreme post-ventral point.

Anatomy. Discussed by Ortmann
(1912a: 355), who regarded this monotypic
subgenus as the most primitive member of
the genus in which the typical features of
the genus are barely indicated.

Plagiola (Truncillopsis) triquetra
(Rafinesque)

Plate 10, figures 1-4

Distribution: Plate 1

Tnnicilld triipictcr Rafinesque 1820, Ann. Gen.
Sci. Physi(iues, Bruxelles 5: 300, pi. 81, figs. 1-4
(chutes de I'Ohio [River, near Louisville, Jef-
ferson Co., Kentucky]; lectotype ANSP 20231
selected by Jolinson and Baker, 1973, Proc.
Acad. Nat. Sci. Phila. 125: 173, pi. 7, fig. 3).

Plagiola from Middle North America • Johnson 249

Unio iriangithris Barnes 1823, American Jour. Sci.
6: 272, pi. 13, fig. 17, a, b (Detroit River
[Michigan]; Bois Blanc Isle [Esse.x Co., On-
tario], figured type Lyceum of Natural History
of New York [destroyed by fire]).

Unio cuneatiis Swainson 1823, Philos. Mag. Jour.,
Edinburgh 61: 112 (no locality; type [lost]).

Unio formosus Lea 1831, Trans. Amer. Philos. Soc.
4: 111, pi. 16, fig. 41 (Ohio River; type not in
the USNM or ANSP [lost]); 1834, Obs. Unio
1: 121.

Unio triangularis pergibosus Gregorio 1914, II Nat.
Siciliano 22: 40, pi. 4, fig. 4 (Sciota [Scioto]
River, Ohio, type presumed to be in Palermo
Mus., Sicily [not seen] ).

Unio triangularis longiusculus Gregorio 1914, II
Nat. Siciliano 22: 40, pi. 4, fig. 5 (Sciota
[Scioto] River, Ohio, type presumed to be in
Palermo Mus., Sicily [not seen] ) .

Truncilla (Truncilla) triquetra ( Rafinesque).
Simpson, 1900, Proc. U. S. Natl. Mus. 22: 517;
1914, Cat. Naiades 1: 6.

Truncilla triquetra (Rafinesque). Ortmann 1909,
Ann. Carnegie Mus. 5: 118; 1912a, Ann. Carne-
gie Mus. 8: 355. Wilson and Clark 1912, U. S.
Bur. Fisheries, Doc. 757: 55; 1914, U. S. Bur.
Fisheries, Doc. 781: 45. Ortmann 1918, Proc.
Amer. Philos. Soc. 57: 585; 1919, Mem. Carne-
gie Mus. 8: 325, pi. 21, figs. 3, 4.

Dtjsnomia {Truncillopsis) triquetra (Rafinesque).
Ortmann and Walker, 1922, Occ. Papers, Mus.
Zool. Univ. Mich., no. 112: 65. Ortmann, 1925,
Amer. Mid. Nat. 9: 359. Frierson, 1927, Check
list N. American naiades, p. 96. Baker, 1928,
Bull. Univ. Wisconsin, no. 1327, p. 296, pi. 86,
figs. 5-7, pi. 70, figs. 4-7. La Rocque, 1967,
Geol. Sur^•. Ohio, Bull. 62 (2): 285, fig. 176.
Haas, 1969, Das Tierreich, pt. 88, p. 479.

Dijsnomia triquetra (Rafinesque). Danglade,
1922, U. S. Bur. Fisheries, Doc. 934, p. 5. Ort-
mann, 1926, Ann. Carnegie Mus. 17: 182. van
der Schahe, 1941, Jour, of Conch. 21: 251. Mor-
rison, 1942, Bur. Amer. Ethnology, no. 129, p.
363. Goodrich and van der Schalie, 1944, Amer.
Mid. Nat. 32: 314. Robertson and Blakeslee,
1948, Bull. Buffalo Soc. Nat. Sci. 19: 112, pi.
11, fig. 9. Murray and Leonard, 1962, Univ.
Kansas, Mus. Nat.' Hist., Misc. Pub. 28, p. 155,
pi. 44, figs. 1-4, text fig. 40. Neel and Allen,
1964, Malacologia 1: 450, fig. 63. Starrett,
1971, Illinois Nat. Hist. Surv. Bull. 30: 340, pi.
4, fig. 21. Clarke, 1973, Mai. Review 6: 64.

Description. Sliell usually of medium
size, reaching up to 80 mm in length. Out-
line long based triangular. Valves slightly
inequilateral, much inflated, solid. Ante-
rior end regularly rounded, posterior end
obliquely truncated. Ventral margin slightly

curved in males, almost straight in females.
Dorsal margin short, almost straight, form-
ing a distinct angle with the obliquely de-
scending posterior margin. Hinge ligament
short. Posterior ridge, high, faintly double,
sharply angled. Dorsal slope very broad,
slightly concave and radially sculptured.
Umbos full and high, turned inward and
anteriorly over a well-marked lunule, lo-
cated slightly anterior of the middle of the
shell; their sculpture is faint consisting of
broken, somewhat doubly-looped ridges.
Surface of the shell smooth except for nu-
merous irregular growth rests. Periostracum
subshiny, tawny to yellow green, with
broken bright green rays and rows of green
mottling.

Left valve with two ragged, subcom-
pressed, triangular, pseudocardinal teeth;
no interdentum. Two short, straight, ele-
vated, granular lateral teeth. Right valve
with two subcompressed, triangular, pseu-
docardinal teeth. One lateral tooth. An-
terior adductor muscle teeth well im-
pressed, posterior ones less so. Pallial line
impressed anteriorly. Umbonal cavities
deep. Nacre white or silvery.

NIale shells grow larger, and are some-
what less sharply triangular than those of
the female.

Female shells have a marsupial swelling
in the area of the posterior ridge consisting
of a slightly ele\'ated, narrowly rounded,
radially sculptured ridge that projects just
below the ventral margin.

Length Height Width
mm mm mm

■90

Little Miami River, Ohio.

Male.

Green River, 8 mi. S

Campbellsville, Taylor Co.,

Kentucky. Male.

As above. Female.

Anatomy and Breeding Season. The
anatomv \\'as discussed by Ortmann
(1912a- 355), who also (1919: 327) de-
termined that the species is bradytictic.

Habitat. Found in riffles \vith stony and
sandv bottoms, in swift currents, usually
deeply buried (Baker, 1928: 298).

250 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Remarks. Plagiola triqiietra ( Rafin-
esque) does not closely resemble any other
member of the genus Plagiola. It may be
distinguished by its long triangular outline,
sharply truncated posterior end and rows
of green mottling. It might be confused
with two other unionids, Triincilla truncata
Rafinesque or Alasmidonta marginata Say,
since both have somewhat superficial re-
semblances to triquctro. T. truncata is more
triangular, with a sharper posterior ridge
and a concave dorsal margin, and A. mar-
ginata lacks distinct pseudocardinal teeth
and has no lateral teeth.

Range. Upper White River system, Mis-
souri; Missouri River drainage, Kansas and
Missouri; Mississippi River system, Wiscon-
sin and Iowa; Illinois River drainage, Illi-
nois; Tennessee and Cumberland River sys-
tems; Green River drainage, Kentucky;
Ohio River system from Indiana to Penn-
sylvania; St. Lawrence River system:
Lakes Michigan and Erie.

Reported in Oklahoma by Simpson
(1914: 6) but not by Isely (1925).

Abundance. This is the most successful
member of the genus in that it is the most
widely distributed and most generally abun-
dant. It occupies more of the formerly
glaciated region than any other Plagiola.

Specimens Examined

White Ri\tr System

Black River Drainage. Missouri: [Black
River], Poplar Bluff, [Butler Co.]. ( MZUM
81269. This important record, based on a
single specimen, has the locality written
on the shell, with the additional data [W.
A.] Marsh, March 3, 1891).

Mississippi River System

Meraniec River Drainage. Missouri:
Bourbeuse River, 5 mi. S Owensville, Gas-
conade Co. (MCZ). Meramec and Big
rivers (Buchanan [collection sites shown
on map, but not listed] ).

Osage River Drainage. Kansas: Marais

des Cygnes River, Ottawa, Franklin Co.
( Scammon ) .

Missouri River Drainage. Kansas: Wa-
karusa River, Lawrence, Douglas Co.
( Scammon ) .

Mississippi River Drainage. Wisconsin:
W^isconsin River, Sauk Co. (MZUM).
Iowa: Mississippi River, Davenport, Scott
Co.; Mississippi River, Muscatine, Musca-
tine Co. {both MCZ).

Illinois River Drainage. Illinois: Kan-
kakee River (Parmalee, 1967). iHinois
River, La Salle, La Salle Co.; Illinois
River, Fulton Co. {both Starrett).

Tennessee Ri\^r System

Powell River Drainage. Tennessee:
Powell River, Shawanee, Claiborne Co.
(CM).

Clinch River Drainage. Virginia: Clinch
River, St. Paul, Wise Co.; Clinch River,
Dungannon and 1.5 mi. below Speers Ferry
bridge, Iwth Scott Co.; CHnch River, 1.5
mi. S Dona, Lee Co. (all MCZ). Tennes-
see: Clinch River below Kyles Ford bridge,
Hancock Co.; Clinch River, 4 mi NW Thorn
Hill, Grainger Co.; Clinch River, CHnton,
Anderson Co.; (all MCZ).

Holston River Drainage. Virginia:
North Fork, Mendota, Washington Co.;
South Fork, Pactolus, Sullivan Co. (both
CM). Tennessee: Holston River, Rogers-
ville, Hawkins Co.; mouth of Holston River,
Austins Grist Mill, Knox Co. (both MCZ).

French Broad River Drainage. Tennes-
see: Nolichuckv River, Chunns Shoals,
Hamblen Co. (CM).

Tennessee River Drainage. Tennessee:
Tennessee River, Knoxville, Knox Co.
(MCZ).

Paint Rock River Drainage. Alabama:
Paint Rock River, Paint Rock, Jackson Co.
(CM).

Flint River Drainage. Alabama: Flint
River, Maysville, Madison Co. (CM).

Elk River Drainage. Tennessee: Elk
River, Fayetteville, Lincoln Co. (MCZ).

Tennessee River Drainage. Alabama:
Tennessee River, Muscle Shoals, between

Plagiola from Middle North America • Johnson 251

Colbert and Lauderdale Cos. (CM). Not
found by Morrison (1942: 363) in the Pick-
wick Basin mounds.

Bear Creek Drainage. Alabama: Bear
Creek, Burleson, Franklin Co. (CM).

Duck River Drainage. Tennessee: Duck
River, \Mlhoite, Marshall Co.; Duck River,
Hardinsons Mill, Maury Co., 12 mi. NW
Lewisburg, Marshall Co.; Duck River, Co-
lumbia, Maury Co. {all MCZ).

Cumberland River System

Cumberland River Drainage. Kentucky:
Beaver Creek, E Rowena Ferry, Russell Co.
(MCZ). Tennessee: Cumberland River,
Goodall Island, Smith Co. (Wilson and
Clark).

Obey River Drainage. Tennessee: Obey
Ri\'er, Duncan Ford, 4 mi. SE Lilydale,
Pickett Co. (MCZ); Obey River, Celina,
Clay Co. (Wilson and Clark).

Cumberland River Drainage. Tennes-
see: Cumberland River, Nashville, David-
son Co. ( MCZ).

Ohio Ri\'er System

\^ abash River Drainage. Illinois: Little
Wabash River, Wayne Co. (MCZ). Indi-
ana: West Fork, \Miite River, Indianapo-
lis, Marion Co.; White River, Rockford,
Jackson Co. (both MCZ). Ohio: Big
Beaver Creek (CM); Wabash River, Re-
covery; both Mercer Co. (MCZ). Indiana:
Salmonia River, Grant Co.; Wabash River,
Lafayette, Tippecanoe Co. {both MCZ).

Green River Drainage. Kentucky: Green
Rixer, S mi. S Campbells\'ille, Taylor Co.;
Green River, Greensburg, Green Co.; Green
River, Rio and Munfordville, both Hart
Co.; Green River, Mammoth Cave, Ed-
mondson Co.; West Fork, Drakes Creek,
Massey Mill, Warren Co.: {all MCZ).

Salt River Drainage. Kentucky: Rolling
Fork, Salt River, Ravwick, Marion Co.
(MCZ).

Ohio River Drainage. Kentucky: Falls
of the Ohio River, near Louisville, Jeffer-
son Co. (Rafinesque). Ohio: Ohio River,
Cincinnati, Hamilton Co. (MCZ).

Miami River Drainage. Ohio: Little
Miami River (MCZ); Miami River (CM).

Scioto River Drainage. Ohio: Olen-
tangy River, Delaware, Delaware Co.; Sci-
oto River, Columbus, Franklin Co.; Big
Darby Creek, 4 mi. S Orient; Scioto River,
Circleville; both Pickaway Co. {all MCZ).

Little Kanawha Ri>er Drainage. West
Virginia: Little Kanawha River, Burnsville,
Braxton Co.; Little Kanawha River, Grants-
ville, Calhoun Co.; North Fork, Hughes
River, Cornwallis, Richie Co. {all CM).

Muskingum River Drainage. Ohio:
Tuscarawas River, New Philadelphia, Tus-
carawas Co.; Mohican River, above conflu-
ence with Kolosing River, Newcastle Twp.,
Coshocton Co. {both MCZ).

Ohio River Drainage. Ohio: Ohio
River, Stubenville, Jefferson Co. (MCZ).

Big Beaver River Drainage. Ohio: Ma-
honing River, near Garretsvalle, Portage
Co. (MCZ). Pennsylvania: Shenango
River, Shenango; Pymatuning, Pymatuning
Township, both Mercer Co. \both CM).

Allegheny River Drainage. Pennsyl-
vania: Leboeuf Creek, Erie Co.; Conneaut
Outlet, Cra\vford Co.; French Creek, Ve-
nango Co. {all CM); Allegheny River,
Kellv and Aladdin, both Armstrong Co.
{both CM).

Monongaliela River Drainage. West
Virginia: West Fork River, Lightburn,
Lewis Co.; West Fork River, Lynch Mines,
Harrison Co. {both CM). Pennsylvania:
Dunkards Creek, Mt. Morris, Greene Co.
(MCZ).

St. Lavi^rence River System

Great Lakes Drainage. (Lake Michi-
gan) Wisconsin: Fox River, Omro, Winne-
bago Co. (Baker). Vlichigan: St. Joseph
River, 2 mi. S Leonidas, St. Joseph Co.
(MZUM): Grand River, Grand Rapids,
Kent Co. (MCZ); Muskegon River, 3 mi.
below Newaygo, Newaygo Co. (MZUM).
(Lake Huron) Michigan: Lake Huron
(Goodrich, 1932). (Lake Erie) On-
tario: Sydenham River, l.S mi. NE Shet-
land, Lamberton Co. (Clarke). Michi-

252 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

gan: Lake St. Clair (Goodrich, 1932).
Ontario: Bois Blanc Isle, Essex Co.
(Barnes); Lake Erie, Rondeau Bay, Kent
Co. (MCZ). Michigan: Huron River, 1 mi.
S Milford, Oakland Co.; Huron River, E
Buck Lake, Livingston Co.; Huron River,
Rockvvood; Lake Erie, La Plaisance Bay;
both Monroe Co. {all MZUM). Ohio:
Auglaize River (MCZ). Swan Creek, To-
ledo, Lucas Co. (CM). Lake Erie, Put-in
Island, Ottawa Co. (Wilson and Clark).
Sandusky River, Fremont, Sandusky Co.
(CM). Grand River, Painsville, Lake Co.
(MCZ). Pennsijlvania: Lake Erie, Presque
Isle Bay, Erie, Erie Co. (CM).

Subgenus Ptagiola s. s. Rafinesque

Type species, Unio inter riiptus Rafinesque
1820; subsequent designation, Herrmann-
sen 1847, 1: 279.

Description. Shell subquadrate, subrhom-
boid, or elliptical somewhat truncated pos-
teriorly. Male smooth on the disk without
any radial furrow. Female with a marsupial
swelling in front of the post basal point;
swelling may or may not extend below tlie
ventral margin, but it is marked by two
distinct sulci.

Key to the Species of Plagiola s. s.

1. Shell not miicli inflated, marsupial swelling

extending below the base 2

Shell greatly inflated, marsupial swelling not
extending below the base arcaeformis

2. Shell moderately thiek, rays generally broken,

sometimes not rayed , 3

Shell very thin and delicate with fine green
rays ._._ lenior

3. Posterior slope acutely angled, from the Mo-

bile-Alabama-Coosa river system _ penita

Posterior slope not acutely angled, from the
Tennessee or Cumberland river systems

-- interrupta

Plagiola {Plagiola) interrupta

(Rafinesque) Plate 10, figures 5-7
Distribution: Plate 7, figure A

Obliquaria {Plagiola) Interrupta Rafinesque 1820,
Ann. Cen. Sci. Physiques, Bnixelles .5: 320 ( le
Kentuky et Ohio [Rivers]. Lectotype ANSP
20257, selected by Johnson and Baker, 1973,

Proc. Acad. Nat. Sci. Phila. 125: 159, pi. 7, fig.
4 and the type locality restricted to the Cumber-
land River).

Unio brevidens Lea 1831, Trans. Amer. Philos. Soc.
4: 75, pi. 6, fig. 6 (Ohio; Cumbedand River,
Tennessee [teste errata sheet]; figured type
[lost]; male and female specimens, subsequently
identified by Lea, USNM 85349). 1834, Obs.
Unio 1 : 85.

Unio intcrruptus (Rafinesque). Conrad, 1834,
New Fresh Water Shells U. S., p. 69; 1838,
Monography Unionidae, no. 10, p. 88, pi. 48.

Truncilla brevidens (Lea). Simpson, 1900, Proc.
U. S. Natl. Mus., 22: 517; 1914, Cat. Naiades
1: 7. Wilson and Clark, 1914, U. S. Bur. Fish-
eries Doc. no. 781: 45.

Truncilla interrupta (Rafinesque). Ortmann, 1918,
Proc. Amer. Philos. Soc. 57: 586.

Dijsnomia {Truncillopsis) brevidens (Lea). Ort-
mann and Walker, 1922, Occ. Papers, Mus.
Zool., Univ. Mich. no. 112, p. 66. Ortmann,
1925, Amer. Midland Nat. 9: 360.

Dijsnomia {Penita) brevidens (Lea). Frierson,
1927, Check list N American naiades, p. 94.
Haas, 1969, Das Tierreich, pt. 88, p. 482.

Dijsnomia brevidens (Lea). Morrison, 1942, Bur.
Amer. Ethnology, Bull. no. 129, p. 363. Neel
and Allen, 1969, Malacologia 1: 448, figs. 59-62.

Description. Shell of medium size, reach-
ing over 80 mm in length, though often not
exceeding 50 mm in length. Outline of male
rhomboid or subtriangular; of female sub-
quadrate. Valves somewhat inequilateral;
males not much inflated; females greatly
inflated, especially old individuals; solid.
Anterior end regularly rounded; posterior
one more broadly rounded. Ventral mar-
gin slightly curved. Dorsal margin of male
straight, forming an obtuse angle with the
obliquely descending posterior margin.
Dorsal margin of female broadly curved
merging imperceptably with the rounded
posterior margin. Hinge ligament promi-
nent. Posterior ridge broadly curved and
faintly double in the male; the posterior
ridge of the female becomes a rather
sharply elevated marsupial swelling,
toothed below and marked with the remains
of former teeth, separated from the rest of
the shell by two distinct sulci. It often
projects well below the base and has a semi-
circular outline on it. Dorsal slope flat,
broad, sometimes with radial sculpture.
Umbos much elevated, elongated, located

Plagiola from Middle North America • Johnson 253

anteriorly, their sculpture consisting of
feeble, double looped bars. Surface of the
disk smooth or clothlike. Periostracum yel-
lowish, tawny, or tawny brown, with nar-
row, broken, radial green rays, sometimes
broken into large dots, especially posterior-
ly.

Left valve with two ragged pseudocardi-
nal teeth, triangular, and of about equal
size; the anterior tooth nan-ow, straight,
directed obhquely forward slightly widen-
ing toward the anterior end; the posterior
tooth triangular; the space between them
triangular and extending to the hinge. In-
terdentum very short and narrow. Two
nearly straight, very short, heavy, obliquely
sculptured lateral teeth. Right valve with
two pseudocardinal teeth, the anterior tooth
small and parallel with the hinge, the pos-
terior tooth long, high, parallel to the an-
terior one, separated from the interdentum
by a deep pit. One well-developed lateral
tooth, often with a parallel vestigial tooth
below it. Umbonal cavities very shallow.
Anterior and posterior adductor muscle
scars well impressed, pallial line distinct.
The marsupial area of the female has a
rounded radial furrow. Nacre white,

Male shells are rhomboid or subtriangu-
lar in outline, moderately inflated, and flat-
tened on the disk.

Female shells are subquadrate, rounded
behind, and greatly inflated. When about
one-third grown the marsupial swelling be-
comes sharply elevated, and decidedly sep-
arated from the rest of the shell by two dis-
tinct sulci.

Length Height Width

70
64

54
49

37
42

Cumberland River, Ten-
nessee. Male.
As above. Female.

Habitat. The species appears to occur in
moderate-sized, clear streams with a rocky
bottom, avoiding the smaller tributaries.
(Wilson and Clark, 1914: 45).

Remarks. Male shells of Plagiola inter-
rupta (Rafinesque) can usually be sepa-
rated from the shells of the other members

of the subgenus Plagiola s. s. by the ten-
dency of the green rays, which are often
present on the entire surface, to be broken
into dots. In P. lenior the rays are confined
to the posterior region and are not broken.
The shell of inter nipta is rhomboid, sub-
triangular or subquadrate in outline,
whereas that of lenior is elliptical; both are
flattened on the disk, but the latter is
smaller, and has a thin, delicate shell. P.
interriipta is easily separated from P. arcae-
formis, as the latter is always much more
inflated, has a stronger posterior ridge, and
a characteristic emarginate posterior mar-
gin.

Female shells of P. interriipta resemble
those of arcaeformis except tlie latter's shells
are much more inflated. The sulci of in-
terrupta are much more acute, and while
its marsupial swelling extends below the
base, the swelling in arcaeformis does not;
instead the base is remarkably flattened.
P. lenior has a marsupial swelling similar to
that of interriipta, but the fonner has such
a thin, delicate, small shell it is unHkely to
be mistaken for the latter.

Old, mature specimens of both sexes of
P. interriipta from the Cumberland River,
and the Holston River of the Tennessee
River system, closely resemble those of
penita of similar maturity from the Mobile-
Alabama-Coosa river system, but they can
be separated morphologically. In general,
the male of interriipta is more rhomboid and
flatter on the disk than is penita, and the
former is covered with green rays which are
broken into dots. In both sexes, if rayed at
all, those of penita are more delicate and
any dots are finer. Females of interriipta
differ from those of penita in that the mar-
supial swelling is marked by two acute
sulci, whereas the posterior one in penita
is obscure or absent, and the dorsal slope
of the latter is much more oblique.

The taxa interriipta and brevidens have
both been used for this species. Say ( 1834:
no. 6 [no pagination]) and Conrad (1834:
69 and 1838, no. 10, p. 88, pi. 48) recog-
nized interriipta Rafinesque; however,

254 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Simpson (19U0a: 517; 1914: 7) did not.
Ortmann and Walker (1922: 66) argued
interrupta could not be recognized from the
original description; Frierson (1927: 79) in-
sisted infcrriipto is Lampsilis- menkiana
(Lea) 1836; and Morrison (1969: 24) as-
serted that 0/;//f/f/^/ria (F/rt^/o/fl) interrupta
Rafinesque 1820 = Unto hrevidem Lea
1831. The present emphasis on the identifi-
cation of the tv'pe as the final criterion in
determining the a\'ailability of a taxon re-
(juires that Rafinesque's name be recog-
nized.

Ranp^e. Tennessee River system, Vir-
ginia, Tennessee and Alabama; Cumberland
River system, Kentucky and Tennessee.

Abundance. The number of specimens in
the collections studied indicate this species
must once have been relatively abundant.
It is considered "threatened" by Stansbery
(1976: 43, 49).

Specimens Examined

Tennessee River System

Powell River Drainage. Virginia: Pow-
ell River, 2.5 mi. S and 7 mi. SW, Jones-
ville, both Lee Co. {both MCZ). Ten-
nessee: Powell River, 8-10 mi. N Tazwell,
Claiborne Co.; Powell River [town of]
Powell River, Campbell Co. {both MCZ).

Clinch River Drainage. Virginia: Clinch
River, Hill Station, 5.5 mi. below Fort
Blackmore, Scott Co.; Station Creek, Lee
Co. {both MCZ). Tennessee: Clinch
River, below Kyles Ford bridge. Clinch
River, Sneedsville, botli Hancock Co. {both
MCZ); Clinch River, Clinton, Clinch
River, Edgmoor, both Anderson Co. {both
MCZ).

Holslon River Drainage. Tennessee:
Holston River, Austins Grist Mill, Knox Co.
(MCZ).

French Broad River Drainage. Ten-
nessee. Nolichucky River (MCZ).

Tennessee River Drainage. Tennessee:
Tennessee River, Knoxville, Knox Co.
(MCZ).

Elk River Drainage. Tennessee: Lower

Elk River (Conrad); Elk River, 4 mi. ESE
Fayetteville, Lincoln Co. (MCZ).

Tennessee River Drainage. Alabama:
Tennessee River, Muscle Shoals, Colbert
and Lauderdale Cos. (CM, MZUM); Ten-
nessee River, Tuscumbia, Colbert Co.
(MCZ); Pickwick Basin, mounds between
Barton, Colbert Co. and Waterloo, Lauder-
dale Co. (Morrison, USNM).

Duck River Drainage. Tennessee: Duck
River, Wilhoite, Marshall Co. (MZUM,
CM); Duck River, Hardinsons Mill, Mau-
ray Co., 12 mi. NW Lewisburg, Marshall
Co. (MCZ); Duck River, Columbia, Mau-
ray Co. (Hinkley and Marsh).

Cumberland River System

Ciiniherland River Drainage. Kentucky:
Cumberland River, Burnside (MCZ); Big
South Fork, opposite Parkers Lake Station
(Wilson and Clark); both Pulaski Co.; Bea-
ver Creek, E Rowena Ferry, Russell Co.
(MCZ).

Canev Fork Drainage. Tennessee: Ca-
ney Fork, Putnam Co. (Wilson and Clark).

Stones River Drainage. Tennessee:
Stones River, Murfreesboro, Rutherford
Co; Stones River, 1.2 mi. W Couchville,
Davidson Co.; (both MCZ).

Cuniherland River Drainage. Tennes-
see: Cumberland River, Nashville, David-
son Co. (USNM).

Ptagiola {Ptagiota) penita (Conrad)
Plate 10, figures 8-15
Distribution: Plate 9, figure B

Unio pcnitiis Conrad 1884, New fresh water shells
United States p. 33, pi. 5, fig. 1 ( Alabama River,
near Claiborne [Monroe Co.], Alabama; figured
holotypeANSP 59860).

Unio metastriatus Conrad 1838, Monography
Unionidac, no. 11 [back cover]; 1840, Ibid., no.
12, p. 104, pi. 57, fig. 2 (Black Warrior River,
Blounts Springs, [Blount Co.], Alabama; figured
types [lost] ).

Unio othcaJoogensis Lea 1857, Proc. Acad. Nat.
Sci. Phila. 9: 32 ( Othcalooga [Oothkalooga]
Creek, Gordon Co., Georgia); 1858, Jour. Acad.
Nat. Sci. Phila. (2) 4: 74, pi. 14, fig. 54,
figured holotype USNM 84615; 1858, Obs. Unio
6:74.

Plagiola from Middle North America • Johnson 255

Unio compactus Lea. 1859, Proc. Acad. Nat. Sci.
Phila. 11: 154 (Etowah River; Conasauga
Ri\er; botli Georgia); 1859, Jour. Acad. Nat.
Sci. Phila. (2) 4: 218, pi. 28, fig. 98, figured
holotvpe USNM 84447 from the former lo-
cahty; 1859, Obs. Unio 7: 36.

Unio modiceUiis Lea 1859, Proc. Acad. Nat. Sci.
Phila. 11: 171 (Conasauga River; Chattanooga
[^Chattooga] River; both Georgia); 1860, Jour.
Acad. Nat. Sci. Phila. (2) 4: 347, pi. 57, fig.
172, figured holotype USNM 84841 from the
former locality; 1860, Obs. Unio 8: 29.

Tnincilla penita (Conrad). Simpson 1900, Proc.
U. S. Natl. Mus. 22: 518; 1914, Cat. Naiades
1: 8.

Dtjsnomia (Penita) penita (Conrad). Frierson,
1927, Check list N American naiades, p. 93;
Haas, 1969, Das Tierreich, pt. 88, p. 481; Hurd,
1974, Ph. D. thesis, p. 97.

Epiohla.wui penita (Conrad). Stansbery, 1976,
Bull. Alabama Mus. Nat. Hist., no. 2: 43, 48, fig.
on p. 49.

Truncilla metastriata (Conrad). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 519; 1914, Cat. Na-
iades 1 : 10.

Dijsnomia (Penita) metastriata (Conrad). Frier-
son, 1927, Check list N American naiades, p.

93. Haas, 1969, Das Tierreich, pt. 88, p. 482.
van der Schalie, 1938, Occ. Papers Mus. Zool.,
Univ. Mich., no. 392, p. 16; 1939, Ibid., no.
407, p. 4; Hurd, 1974, Ph. D. thesis, p. 95.

Tnincilla othcaloogensis (Lea). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 521; 1914, Cat. Na-
iades 1: 17.

Dtjsnomia (Penita) othcaloogensis (Lea). Frier-
son, 1914, Check hst N American naiades, p.

94. Haas, 1969, Das Tierreich, pt. 88, p. 484.
van der Schalie, 1938, Occ. Papers, Mus. Zool.,
Univ. Michigan, no. 392, p. 16. Hurd, 1974,
Ph. D. tliesis, p. 96.

Truncilla cornpacta (Lea). Simpson, 1900, Proc.

U. S. Natl. Mus. 22: 518; 1914, Cat. Naiades 1:

9.
Truncilla modicella (Lea). Simpson, 1900, Proc.

U. S. Nad. Mus. 22: 518; 1914, Cat. Naiades 1:

11.
Dijsnomia (Penita) modicella (Lea). 1927, Check

list N American naiades, p. 94. Haas, 1969, Das

Tierreich, pt. 88, p. 484.

Description. Shell of medium size reach-
ing 60 mm in length. Outline of male rhom-
boid or subtriangular, of female subrhom-
boid or quadrate. Valves somewhat
inequilateral, males usually moderately in-
flated, females considerably inflated, solid.
Anterior end regularly rounded, posterior
end more broadly rounded. Ventral margin
sli^htlv curved. Dorsal margin of male

straight, forming an obtuse angle with the
obliquely descending posterior margin.
Dorsal margin of female broadly curved
merging imperceptably with the rounded
posterior margin. Hinge ligament promi-
nent. Posterior ridge broadly curved, some-
times quite imperceptable in the male.
When about one-half grown, the posterior
ridge of the female often becomes a rather
sharply-elevated, narrow, rounded marsu-
pial swelling, often slightly toothed below,
and sometimes marked with the remains of
former teeth, separated from the anterior of
the shell by a sulcus. The marsupial swell-
ing often projects below the base line, and
when it does is rather long and has a semi-
circular outline on it. Dorsal slope flat and
narrow, sometimes with radial sculpture.
Umbos much elevated, located anteriorly,
their sculpture not observed. Surface of the
disk smooth. Periostracum yellowish,
tawny, or tawny brown, often with broken
radial green lines, sometimes broken into
inconspicuous dots, arrowhead markings or
darker color arranged radially on the pos-
terior part.

Left valve with two ragged pseudocardi-
nal teeth, triangular and of about equal
size; the anterior tooth is narrow, straight,
directed obliquely forward, slightly widen-
ing toward the anterior end; the posterior
tooth is triangular, the space between them
is triangular and extending to the hinge.
Interdentum very short and narrow. Two
nearly straight, short, heavy, obliquely
sculptured lateral teeth. Right valve with
two pseudocardinal teeth, the anterior tooth
small, parallel to the hinge, the posterior
tooth long, high, parallel to the anterior one,
separated from the interdentum by a deep
pit. One well-developed lateral tooth often
with a parallel vestigial tooth below. Um-
bonal cavities very shallow. Anterior and
posterior muscle scars well impressed, pal-
lial line distinct. The marsupial area of the
female showing a rounded radial furrow.
Nacre white, or bluish white.

Male shells are rhomboid or subtriangu-
lar in outline, and moderately inflated.

256 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Female shells are subrhonil^oid or ({iiad-
rate, subtruncated beliind and often
greatly inflated. When about one-third
grown the marsupial area becomes swollen
and elevated, separated from the anterior
part of the shell by a distinct sulcus.

LenKth Height Width
mm mm mm

Conasauga River, 4.3 mi.
W Eton, Murray Co.,
Georgia. Male.

11.5

As above. Female.

Alabama River, Selma,
Autauga Co., Alabama.
Male.

As above. Female.

Remarks. Phi<i,iola penita (Conrad) of
the Mobile-Alabama-Coosa river system re-
sembles P. internipta of the Tennessee and
Cumberland river systems. Males of the
former are, in general, more quadrate, or
proportionally higher than those of inter-
nipta. P. penita remain quadrate, or sub-
triangular, throughout their range though
specimens from the upper reaches of the
Coosa River drainage do not grow very
large and are often rayless. P. internipta
from the upper reaches of headwater
streams become rather more rhomboid and
flattened on the disk than specimens from
elsewhere, though still growing up to 50
mm in length and exhibiting distinct green
rays broken into dots.

Females of P. penita and internipta, from
those habitats where each reach their maxi-
mum size, rather closely resemble one an-
other; the marsupial swelling of both is
marked from the di.'k by an anterior sul-
cus, while the latter also has a sharp pos-
terior sulcus that is slight, or lacking, in
penita. P. penita has a posterior margin
that slopes more obliquely than that of in-
ternipta; this renders the dorsal slope of the
former much narrower. Females of penita
from the upper reaches of the Coosa River
drainage are usually small, rayless, and ap-
pear barely to reach sexual maturity. These
were recognized as othcaloo<^emis by Hurd
(1974: 96) who followed the usage of
authors since Simpson (1914: 8). The lo-
calities of the lots, included under this taxon

by Hurd, in the Museum of Comparative
Zoology, are with their catalogue numbers,
since the latter are the only data given by
him. (See under Specimens Examined.)
Simpson (1914: 9) recognized compactus
(included with modiceUus under otlicaloo-
gemi'i by Hurd), and noted that female
shells are generally, though not always,
shorter than those of penita; that they are
rather narrower and more compressed in
front, and that the posterior end is some-
what evenly rounded instead of being
obliquely truncate. Simpson (1914: 10)
suggested that U. metastriatws Conrad was
nothing more than a variety of compactus,
in which case the later taxon would have
priority. The former was recognized as a
valid species by Hurd (1974: 95). Stans-
bery (1976: 49) recognized both metastri-
ata and penita as valid and said of the lat-
ter, "This species is very similar to, yet
distinct from EpiohJasma nietastriata,"
without further elucidation.

The female shells of penita are variable
as to the extent of the marsupial swelling.
Variation in the amount of inflation of the
valves occurs in both sexes. Specimens
from the larger rivers tend to attain greater
size, are more inflated and are more often
rayed than those occurring in smaller head-
water streams. In the latter, shells are
small, not greatly inflated, almost always
rayless and with a smooth yellow perio-
stracum. When rayed at all, those of
penita are narrower and more delicate than
those of internipta from the Tennessee and
Cumberland river systeins. When the rays
of the former are broken into dots, thev are
much finer than those of internipta.

Van der Schalie (1938a: 27) noted that
no Dymomia [Pkiiiiola] occur in the river
systems on either side of the Mobile-Ala-
bama-Coosa river system and correctly sug-
gested that it, and a number of other gen-
era, arrived there through a former
confluence with the Tennessee River sys-
tem.

Range. Mobile-Alabama-Coosa river sys-
tem, Georgia, Alabama, and Mississippi.

Plagiola from Middle North America • Johnson 257

Abundance. The number of specimens
in the collections studied indicate this spe-
cies must have been relatively abundant.
Hurd (1974: 170-180), during 1971-73,
made an extensive survey of the Coosa
River drainage for Unionidae. He col-
lected at 194 stations but found onlv 28
specimens of Plagiola from four localities,
all of which he regarded as otiwaloogensis
(pp. 42, 96). Stan^sbery (1976: 43, 48, 49)
lists othcaloogensis and penita as having an
"endangered status" and metastriata as hav-
ing a "threatened status."

Specimens Examined
Mobile-Alabama-Coosa River System

Conasauga River Drainage. Georgia:
Conasauga River, Beaverdale, Whitfield
Co. (MCZ 200353); Conasauga River, Up-
per Kings Bridge (MCZ 933788); Cona-
sauga River, Lower Kings Bridge (MCZ);
Conasauga River, 4.3 mi. W Eton ( MCZ
36620 & 214237); all Murray Co. Cona-
sauga River, Fikes Ford, 1.4 mi. N Resaca,
Gordon Co. (MCZ).

Oothkalooga Creek Drainage. Georgia:
Oothkalooga Creek, Gordon Co. (Lea,
USNM).

Oostanaula River Drainage. Georgia:
Oostanaula River, Rome, Floyd Co. (MCZ).

Etowah River Drainage. Georgia: Eto-
wah River, Rome, Floyd Co. (MCZ).

Chattooga River Drainage. Georgia:
Chattooga River (MCZ 16348, 16532, 28711,
28809).

Coosa River Drainage. Alabama: Mill
Creek (MCZ 51518, 16444); Coosa River,
Stackland (MCZ); both Cherokee Co.
Coosa River, Gadsden, Etowah Co. ( MCZ ) .
Coosa River, old lock 5, 6 mi. SW, Lincoln;
Coosa River, Fort William Shoals; both
Talladega Co. Coosa River, Weduska
Shoals and Three Island Shoals, both
Shelby Co. Coosa River, Wetumpka, El-
more Co. (all MCZ).

Cahaha River Drainage. Alabama:
Cahaba River, Henry Ellen, Lovick, Grants
Mill, and E. Merkel, all Jefferson Co. (all

MZUM); Buck Creek, Helena (MCZ,
MZUM); Cahaba River, Nunley Ford
(MZUM); both Shelby Co.; Cahaba River,
Lily Shoals (MCZ), and 10 mi. above Cen-
terville (MZUM), both Bibb Co.; Cahaba
River, 8 mi. N Sprott, and 5 mi. NE Mar-
ion, both Perry Co. (both MZUM).

Alahania River Drainage. Alabama:
Alabama River, Selma, Dallas Co. (MCZ);
Alabama River, near Claiborne, Monroe Co.
( Conrad ) .

Black Warrior River Drainage. Ala-
bama: Black Warrior River, Blounts
Springs, Blount Co. ( Conrad ) ; Black War-
rior River, Squaw Shoals, Jefferson Co.;
Black Warrior River, Tuscaloosa Co. ( both
MCZ);

Tonihighee River Drainage. Missis-
sippi: East Fork Tombigbee River, 3 mi.
W Smithville, Monroe Co. (OSM, MCZ).
Alabama: Tombigbee River, Epes, Sum-
ter Co. (MZUM).

Plagiola {Plagiola) arcaeformis (Lea)
Plate 11, figures 1-4
Distribution: Plate 8, figure B

Unto arcaeformis Lea 1831, Trans. Amer. Philos.
Soc. 4: 116, pi. 17, fig. 44 (Tennessee River;
figured type [lost], male and female specimens
subsequently identified by Lea USNM 84422);
1884, Obs. i : 126.

Unio nexus Say 1831, Transylvania Jour. Medicine
4: 527 (Cumberland River, Nashville [David-
son Co.], Tennessee); 1834, Amer. Conch., no.
6, pi. 51, figured type [lost].

TrunciUa arcaeformis (Lea). Simpson, 1900, Proc.
U. S. Natl. Mus. 22: 519; 1914, Cat. Naiades 1:
12. Wilson and Clark, 1914, U. S. Bur. Fisher-
ies Doc. 781, p. 46. Ortmann, 1918, Proc. Amer.
Philos. Soc. 57: 586.

Dysnomia (TriinciUopsis) arcaeformis (Lea). Ort-
mann, 1925, Amer. Mid. Nat. 9: 359.

Dysnomia (Penita) arcaeformis (Lea). Frierson,
1927, Check list N American naiades, p. 94.
Haas, 1969, Das Tierreich, pt. 88, p. 482.

Dysnomia arcaeformis (Lea). Morrison, 1942,
Bur. Amer. Ethnology Bull. 129, p. 363. Stans-
bery, 1970, Malacologia 10: 19, pi. 1, figs. 5, 6;
1971, Svmposium of rare and endangered Moll.
U. S., p.' 18a, figs. 1-2.

Description. Shell of medium size, reach-
ing 70 mm in length. Outline of shell sub-

258 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

(jiiadrate or subrhomboid. Valves inequi-
lateral, greatly inflated, females especially
so, solid. Anterior end regularly rounded;
posterior end more broadly rounded. Ven-
tral margin slightly cur\'ed in males, with
a slight emargination before the posterior
ridge; ({uite straight in females. Dorsal
margin straight forming an obtuse angle
with the obliquely descending, emarginate,
posterior margin. Hinge ligament promi-
nent. Posterior ridge full, high, double or
triple, subangulate, subtruncate behind the
ridge. \Mien about half grown the posterior
ridge of the female becomes a marsupial
swelling, slightly toothed below, marked
with faint tooth sculpture at rest lines and
with faint radial sculpture, separated from
the rest of the shell by two distinct but
broad sulci. Marsupium much flattened
below, does not project below the base
which is remarkably flattened. On the dor-
sal slope above the upper posterior ridge
is usually a shallow radial furrow. Umbos
full, much elevated, almost touching one
another, located anteriorly, their sculpture
consisting of undulating ridges. Surface of
the shell smooth, or cloth-like. Periostra-
cum tawny to yellowish-green, usually with
delicate green rays over the entire surface.
Left valve with two ragged pseudocardi-
nal teeth, triangular, and of about equal
size, the space between them triangular and
extending to the hinge. Interdentum very
short and narrow. Two nearly straight, very
short, heavy, obliquely sculptured lateral
teeth. Right valve with one large pseudo-
cardinal tooth, sometimes with a smaller
tooth before or behind it. When present,
the anterior tooth is small and parallel with
the hinge. The pit before the interdentum
is deep. One well-developed lateral tooth,
sometimes with a vestigial tooth below.
Umbonal cavities moderately deep. An-
terior and posterior adductor muscle scars
well impressed. Pallial line distinct. The
marsupial area of the female is much thin-
ner than the surrounding shell and has a
rounded radial furrow. Nacre white.

Length Height Width
mm mm mm

Cumberland River, Ten-
nessee. Male.

Tennessee River, Tennes-
see. Female.

Remarks. Plagiola arcaefonnis (Lea)
may be distinguished from any other mem-
ber of the genus by the extreme inflation
of both the male and female shells, by the
radial furrow above the posterior ridge
and by the emarginate posterior margin.

Both male and female shells are con-
siderably alike in outHne, though the lat-
ter are less elevated and more inflated.
Old females are extraordinarily inflated.
The marsupial swelling is considerably in-
flated toward the base, but is remarkably
flattened so that it scarcely projects below
the base.

Young males of arcaeformis might pos-
sibly be mistaken for P. turgidula, except
the latter is not as high or inflated and has
a shallow radial furrow below rather than
above the posterior ridge.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem, Kentucky and Tennessee.

Abundance. Generally not found in
great numbers, but reported to be locally
abundant in the Holston River drainage of
the Tennessee River system by Ortmann
(1918: 586). "The entire range of this
species is now under a series of impound-
ments. It has not been collected in over
half a centuiy and hence is presumed ex-
tinct," (Stansbery, 1976: 43, 50).

Specimens Examined
Tennessee River System

Clintli River Drainage. Tennessee:
Clinch River, Clinch River Station, Clai-
borne Co. (CM); Clinch River, Oakman,
Grainger Co. (CM); Clinch River, Clinton,
Anderson Co. (MCZ).

Holston River Drainage. Tennessee:
Holston River, near Rogersville (MCZ);

Placiola from Middle North America • Johnson 259

Holston River, Austin Mill (CM), both
Hawkins Co.; Holston River, 4 mi. above
Morristown, Hamblen Co. (MZUM); Hol-
ston River, Holston Station; Holston River,
Turley Mill, Noeton; botJi Grainger Co.
(both CM). Holston River, Cant Island,
nr. Straw Plains, Jefferson Co. (MZUM);
mouth of Holston River, Austins Grist Mill;
Holston River, Knoxville; botli Knox Co.
{both MCZ).

French Broad River Drainage. Tennes-
see: French Broad River, Boyd Creek,
Sevier Co. (CM).

Tennessee River Drainage. Tennessee:
Tennessee River, Knoxville, Knox Co.
(MCZ). Alabama: Tennessee River,
Bridgeport, Jackson Co. (MCZ); Tennessee
River, Florence, Lauderdale Co. (MCZ);
Tennessee River, Tuscumbia, Colbert Co.
(MCZ); Pickwick Basin, mounds between
Barton, Colbert Co. and Waterloo, Lau-
derdale Co. (Morrison, USNM).

Cumberland River System

Cumberland River Drainage. Ken-
tucky: Cumberland River (MCZ); Big
South Fork of Cumberland River, 2 mi.
above Burnside, Pulaski Co. (Wilson and
Clark). Tennessee: Cumberland River,
Nashville, Davidson Co. (Say).

Plagiola {Plagiola) lenior (Lea)
Plate 11, figures 5-6
Distribution: Plate 7, figure C

Unio lenis Lea 1840, Proc. Anier. Philos. Soc. 1:

286 (Stones River, Tennessee), non Conrad

1840. Changed to:
Unio lenior Lea 1842, Trans. Amer. Philos. Soc.

8: 204, pi. 12, fig. 18, figured holotype USNM

86130; 1842, Obs. Unio 3: 42.
Tnincilla lenior (Lea). Simpson, 1900, Proc. U. S.

Natl. Mus. 22: 518; 1914, Cat. Naiades 1: 11.

Ortniann, 1918, Proc. Amcr. Philos. Soc. 57:

587.
Dijsnomia (TninciUopsis) lenior (Lea). Ortmann,

1924, Amer. Mid. Nat. 9: 34; 1925, Amer. Mid.

Nat. 9: 360.
Dijsnomia (Penita) lenior (Lea). Frierson, 1927,

Check list N American naiades, p. 94. Haas,

1969, Das Tierreich, pt. 88, p. 484.

Dysnomia lenior (Lea). Stansbery 1970, Mala-
cologia 10: 19, pi. 2, figs. 3, 4; 1971, Sympcsium
of rare and endangered moll. U. S. p. 18f, figs.

3, 4.

Description. Shell usually of small size,
reaching almost 40 mm in length. Outline
of male subelliptical; of female subquad-
rate, a little narrower anteriorly. \'alves
inequilateral, subinflated to inflated, thin
and delicate. Anterior end regularly
rounded, posterior end of male slightly more
broadly rounded; subtruncate in females.
Ventral margin slightly curved. Dorsal
margin almost straight, forming a slight an-
gle with the obliquely descending posterior
margin. Posterior ridge triple. Dorsal slope
slightly rounded. Below the third ridge in
females is a radially sculptured, toothed,
raised marsupial swelling separated from
the rest of the shell by two sulci, the more
posterior one acute, ending in an emargina-
tion below the middle of the posterior
slope. The marsupium projects slightly be-
low the base and has a semicircular outline
that extends posteriorly beyond it. Umbos
slightly elevated, located toward the an-
terior third of the shell, their sculpture con-
sisting of doubly-looped bars. Surface of
the shell smooth with delicate growth lines.
Periostracum subshiny, pale ashy-greenish,
or yellowish green, with numerous, narrow
green rays posteriorly.

Left valve with two delicate, subcom-
pressed, pseudocardinal teeth; no interden-
tum. Two short, delicate, lateral teeth.
Right vaKe with one pseudocardinal and
one lateral tooth. Umbonal cavities very
shallow. Anterior adductor muscle scars
well impressed, posterior ones less so. Pal-
lial line faint. The shell is thin, especially
so in the marsupial furrow of the female.
Nacre bluish white.

Length Height Width
mm mm mm

38 23 15

Eastern Tennessee. Male.

Stones River, Tennessee.
Female. Identified by
Lea.

Remarks. Plagiola lenior (Lea) is dis-

260 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

tingiiished from other members of Plagiola
by its thin shell and delicate green rays,
which are restricted to the posterior end.
The male might be mistaken for a young,
pale colored Villo.sa nehulosa ( Conrad ) , ex-
cept that it has fine denticles on the posteri-
or margin. The female has, in addition to
the delicate shell, an acute sulcus behind the
marsupial swelling that ends in an emargi-
nation below the middle of the posterior
slope.

Range. Tennessee River system, Virginia,
Tennessee and Alabama. Cumberland
River system, restricted to Stones River,
Tennessee.

Abundance. A rare species that appears
to be somewhat locally abundant in the
Paint Rock River, Alabama (Ortmann,
1918: 587). "The last known population
of this species is now covered by the Priest
Reservoir on the Stones River in Tennes-
see," (Stansbery, 1970: 19). It was last
collected in the Stones River by Stansbery
and Jenkinsen in 1965. Considered to be ex-
tinct in the Clinch River, above Norris
Dam, Tazewell, Clairbone Co., Tennessee
(Stansberv, 1972: 22). Listed as "extir-
pated" by Stansbery (1976: 43.50).

Specimens Examined
Tennessee River System

Powell River Drainage. Virginia: Pow-
ell River, Dickson Ford, 3 mi. SE Jones-
ville, LeeCo. (MZUM).

Clinch River Drainage. Virginia:
Clinch River, Speers Ferry, ( CM ) ; Clinch
River, Maness ( MZUM ) ; both Scott Co.

Holston River Drainage. Tennessee:
North Fork, Holston River, Rotherwood,
Hawkins Co. (CM); South Fork, Holston
River, Pactolus, Sullivan Co. (CM); Hol-
ston River, Church Hill, Hawkins Co.
(MCZ); Holston River nr. Knoxville, Knox
Co. (Andrews, Walker colln. UZUM).

Paint Rock River Drainage. Tennessee:
Paint Rock River, Holly Tree, Trenton,
Paint Rock, all Jackson Co. {all CM); Paint

Rock River, Woodville Jackson Co.
(USNM, MZUM).

Duck River Drainage. Tennessee: Duck
River, Columbia, Maury Co. (Hinkley and
Marsh ) .

Cumberland River System

Stones River Drainage. Tennessee:
Cumberland River ( [written on shell]
USNM) Stones River, 1.2 mi. W Couch-
ville, Davidson Co. (MCZ).

Subgenus Torutosa Frierson

Torulosa Frierson 1927, Check list N American
naiades, pp. 11, 94. Type species, Amblema
torulosa Rafinesque, original designation, teste
Errata et Corrigenda.

Capsacformis Frierson 1927, Check list N Ameri-
can naiades, pp. 11, 95. Type species, Unio
capsacformis Lea, original designation, teste Er-
rata et Corrigenda.

Description. Shell usually ovate, obovate
or elliptic.il, occasionally triangular, male
shell w-th a wide radiating depression, of
varying depth, in front of the posterior
ridge. The depression usually ends in an
emargination. Female with a thin, com-
pressed or subcompressed marsupial swell-
ing that occupies the entire post-basal re-
gion.

Anatomy. In his discussion of the anato-
my of rangiana ( = torulosa), florentina and
capsacformis, Ortmann (1912a: 358-360)
mentions no differences that would indicate
any of these species should be placed in
different subgenera.

Discussion. Frierson (1927) did not give
descriptions of any of his new subgenera,
but merely cited type species. He arbi-
trarily created subgenera for every assem-
blage of four or five species in genera of
any size. Of the species recognized in this
paper that would fall into Capsaeformis —
namely capsaeformis, florentina and tur-
gidula — all that can be said to differentiate
them from other Torulosa is that they have
thinner, smaller shells and that the radial
depression in front of the posterior ridge in

Plagiola from Middle North America • joJtnson 261

capsaeforrnis and florentina is often faint.
These minor conchological characters do
not warrant the use of Capsaeforrnis as a
subgenus.

Key to the Species of Torulosa

1. Shell elliptical, ovate, or obovate 2

Shell subtriangular, subrhomboid, subquad-
rate or trapezoid 5

2. Shell thin, small, with a smooth surface 3

Shell thick, often large, generally with some
nodules torulosa

3. Surface light yellow, honey colored, or yel-
lowish green 4

Surface green or greenish brown .. capsaeforrnis

4. Rays distributed evenly over the entire sur-
face - ttirgidula

Rays somewhat darker in the radial depres-
sion florentina

5. Radial furrow oblique 6

Radial furrow subxertical biemarginata

6. Furrow wide and shallow, periostracum

shiny sampsoni

Furrow narrow and deep, periostracum sub-
shiny propinqua

Plagiola {Torulosa) torulosa

(Rafinesque)

Plate 11, figures 7-11
Plate 12, figures 1-5
Distribution: Plate 9, figure A

Amblema torulosa Rafinesque 1820, Ann. Gen.
Sci. Physiques, Bruxelles 5: 314, pi. 82, figs.
11, 12 (I'Ohio et le Kentuky [Rivers]; holotype
ANSP 20218 from the Kentucky River, re-
figured bv Johnson and Baker, 1973, Proc. Acad.
\at. Sci. Phila. 12.5: 173, pi. 7, fig. 5).

Amblema torulosa angulata Rafinesque 1820, Ann.
Gen. Sci. Physiques, Bruxelles 5: 315 (I'Ohio et
le Kentuky [Rivers]; type [lost]).

Amblema gibbosa Rafinesque 1820, Ann. Gen. Sci.
Physiques, Bnixelles 5: 315 (I'Ohio et les rivi-
eres adjacentes; lectotype ANSP 20232 from the
Ohio River, selected by Johnson and Baker,
1973, Proc. Acad. Nat. Sci. Phila. 125: 156, pi.
7, fig. 6) [also the varieties: olivacea, radiata
and difformis; all t>'pes lost].

Unto perplexus Lea, 1831, Trans. Amer. Philos.
Soc. 4: 112, pi. 17, fig. 42 (Ohio River [Cincin-
nati, Hamilton Co., Ohio]), figured holotype
USNM 84.324; 1834, Obs. Unio 1: 122.

Unio gibbosus pcrobliquus Conrad 1836, Monog-
raphy Unionidae, no. 6, p. 51, pi. 27, fig. 2 (\N'a-
bash River, Indiana; Detroit River, Michigan,
type lost). The prominent knobs on the mid-
dle of the shell suggest the figured specimen

came from the former locality and not from the
Great Lakes drainage, where knobs are usually
lacking. The type locality is here restricted to
the Waba.sh River, Indiana.
Unio perobliquus Conrad, 1837, Monography

Unionidae, no. 8 [back cover].
Vnio rangianus Lea 1838, Trans. Amer. Philos. Soc.
6: 95, pi. 18, fig. 56 (Ohio River, near Cincin-
nati [Hamilton Co.]; Mahoning River, near Po-
land [Mahoning Co.]; both Ohio); type [lost].
Lectotvpe, here selected, USNM 84798 (pi.
12, Fig. 3), USNM 84798 from the latter
locality; 1838, Obs. Unio 2: 95.
Unio cincinnaticnsis Lea 1840, Proc. Amer. Philos.
Soc. 1: 285 (Ohio River, Cincinnati [Hamil-
ton Co.], Ohio); 1842, Trans. Amer. Philos. Soc.
8: 194, pi. 8, fig. 4, figured holotype USNM
84199; 1842, Obs. Unio. 3: 32.
Unio obliquus Potiez and Michaud 1844, Galerie
Moll. Cat. Mus. Douai 2: 153, pi. 48, figs. 3, 4
(I'Ohio [River]); figured type in Mus Douai
[not seen], non Lamarck 1820.
Unio phillipsii Reeve 1864, Conch. Iconica 16,
Unio, pi. 4, species 15 (North America), figured
holotype BMNH 196481, non Conrad 1835.
Unio gubernaculum Reeve, 1865, Conch. Iconica
16, Unio, pi. 28, species 146 (Hab.?), figured
holotype BMNH 1965203.
Truncilla (Scalenaria) sulcata delicata Simpson
1900, Proc. U. S. Natl. Mus. 22: 520 (Detroit
River, [Amherstburg, Essex Co., Ontario]; holo-
type USNM 160853 [only specimen]); 1914.
Cat. iNaiades 1: 16. Approaches rangiana (Lea),
teste.
Dijsnomia sulcata delicata Simpson Momson.
1942, Bur. Amer. Ethnology Bull. no. 129, p.
364. La Rocque, 1967, Geol. Survey Ohio, Bull,
no. 62 (2): 280.
Truncilla (Pilea) perplexa (Lea). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 522; 1914, Cat. Na-
iades 1 : 24.
Dijsnomia perplexa (Lea). Goodrich and van der

Schalie, 1944, Amer. Mid. Nat. 32: 314.

Truncilla torulosa (Rafinesque). Ortmann, 1918.

Proc. Amer. Philos. Soc. 57: 589. Ball, 1922,

Ecology 3: 115.

Dijsnomia (Pilea) torulosa (Rafinesque). Ortmann

and Walker, 1922, Occ. Papers, Mus. Zool.,

Univ. Mich., no. 112, p. 69. Ortmann, 1925,

Amer. Mid. Nat. 9: 363. La Rocque, 1967,

Geol. Survey Ohio. Bull. 62 (2): 280, fig. 173.

Dijsnomia torulosa (Rafinesque). Ortmann, 1926,

Ann. Carnegie Mus. 17: 182. Morrison, 1942,

Bur. Amer. Ethnology, Bull. 129, p. 364. Stans-

bery 1970, Malacologia 10: 20.

Dijsnomia (Torulosa) torulosa (Rafinesque).

Frierson, 1927, Check list N American naiades,

p. 94. Haas, 1969, Das Ticrreich, pt. 88, p. 485.

Truncilla (Pilea) perplexa rangiana (Lea). Simp-

262 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

son, 1900, Proc. U. S. Natl. Mus. 22: 523. Ort-
mann, 1909, Ann. Carnegie Mus. 5: 188. Simp-
son, 1914, Cat. Naiades 1: 25.

Trunctlla rangiana (Lea). Ortmann, 1912, Ann.
Carnegie Mus. 8: 358, fig. 28 (anatomy); 1919,
Mem. Carnegie Mus. 9: 331, pi. 21, figs. 5-7.
Ball, 1922, Ecology 3: 117. Danglade, 1922,
U. S. Bur. Fisheries. Doc. no. 934: 5.

Dijsnomia {Torulosa) rangiana (Lea). Frierson,
1927, Check list N American naiades, p. 95.

Dijsnomia pcrplcxa rangiana (Lea). Goodrich,
1932, Moll. Mich. p. 113. van der Schalie, 1941,
Jour. Conch. 21: 251.

Dii-snomia (Pilca) torulosa rangiana (Lea). La
Rocque, 1967, Geol. Survey Ohio, Bull. 62 (2):
283.

Dijsnomia {Torulosa) torulosa rangiana (Lea).
Haas, 1969, Das Tierreicli, pt. 88, p. 486.

Truncilla (Pilea) perplexa cincinnatiensis (Lea).
Simpson, 1900, Proc. U. S. Natl. Mus. 22: 523;
1914, Cat. Naiades 1: 26.

Dijsnomia (Torulosa) torulosa cincinnatiensis
(Lea). Frierson, 1927, Check list N American
naiades, p. 94. Haas, 1969, Das Tierreich, pt.
88, p. 486.

Dijsnomia torulosa cincinnatiensis (Lea). Mor-
rison, 1942, Bur. Amer. Ethnology, Bull. no.
129: 365.

Dijsnomia (Pilea) torulosa cincinnatiensis (Lea).
La Rocque, 1967, Geol. Survey Ohio, Bull. 62
(2): 283.

Truncilla torulosa giibemaculum (Reeve). Ort-
mann, 1918, Proc. Amer. Philos. Soc. 57: 590.
Ball, 1922, Ecology 3: 114.

Dijsnomia torulosa guhernaculum (Reeve). Ort-
mann, 1926, Ann. Carnegie Mus. 17: 182.
Stansbery, 1971. Symposium of rare and en-
dangered moll. U. S., p. 18f, figs. 49, 50.

Dijsnomia (Torulosa) torulosa guhernaculum
(Reeve). Frierson, 1927, Check list N American
naiades, p. 95.

Epioblasma torulosa guhernaculum (Reeve).
Stansbery, 1972, Amer. Mai. Union, Bull, for
1972, p. 22.

Description. Shell of medium size,
reaching almost 90 mm in length. Outline
irregularly ovate, elliptical or obovate.
X^ilves inequilateral, sul)inflated to inflated,
solid. Anterior end regularly rounded, pos-
terior end of male slightly produced; more
broadly rounded in females. Ventral mar-
gin slightly curved. Dorsal margin curved
forming an indistinct angle with the
obliquely descending posterior margin.
Hinge ligament short. Posterior ridge of the
male rather low, narrowly rounded, sepa-

rated from a similar medial ridge by a
broad fvurow of varying depth that ends
in an emargination between the somewhat
produced ridges. Both the ridges and the
furrow vary from being smooth to having
elevated knobs. The marsupial swelling in
the female is sometimes marked by a num-
ber of small radial furrows, but the ridges
become obscure. The rounded marsupial
swelling extends from the middle of the
base to the upper part of the posterior end.
Umbos full, somewhat turned over a small
lunule, located toward the anterior third of
the shell, their sculpture feeble and cor-
rugated. Surface of the sliell with many
distinct growth lines. Periostracum smooth
and shiny, tawny, yellowish green, or straw
colored, usually with numerous green rays.

Left valve with two triangular pseudo-
cardinal teeth; slight interdentum. Two
long, almost straight, lateral teeth. Right
valve with one large pseudocardinal tooth
with a smaller tooth before it. One lateral
tooth, sometimes with a vestigial tooth be-
low. Umbonal cavity shallow. Anterior and
posterior adductor muscle scars and pal-
lial line, well impressed. The shell is thin
in the marsupial region of the female espe-
cially toward the margin. Nacre white to
salmon-red.

Male shells are generally irregularly ovate
with a rather wide radial furrow of varying
depth ending in a broad sinus, often bluntly
pointed behind.

Female shells are generally obovate,
larger than the male, having a large, flat-
tened, rounded marsupial swelling extend-
ing from the middle of the base to near the
upper part of the posterior end, which is
thin and often a darker green than the rest
of the shell.

Length Height Width
mm mm mm

68 57 42 Wabash River, Indiana.

Male.

85 59 38

As above. Female.

IIaI)itat. Lives in coarse sand and gravel,
in current, and in water from a few inches
to 4 to 6 feet (Parmalee:1967: 62).

Plagiola from Middle North Americ^. • Joh.r^jon 263

Remarks. Plagiola torulosa (Rafinesque)
exhibits considerable ecophenotypic varia-
tion as to sculpture, or the lack of it, and
to obesity. Ball (1922: 116) convincingly
showed the relationship in this species be-
tween strong tubercle development and
large stream-flow and between the lack of
tubercles and small stream flow, as well as
toridosa's tendency to be more compressed
in smaller streams. Ortmann (1918: 590)
had previously noted that in the Tennessee
River, "the typical torulosa has a radial
row of prominent knobs across the middle
of the shell. But these knobs vary greatly,
and in the upstream direction, they have a
tendency to become reduced, finally disap-
pearing, thus passing into the condition seen
in the next form [guhernaculum Reeve].
From the typical torulosa, this variety dif-
fers by the poorly developed, or wanting,
knobs, and by the rather more compressed
shell. This is the headwaters form of toru-
losa and begins to take place in the [Ten-
nessee River] in the vicinity of Knoxville
[Knox Co., Tennessee]." Ortmann further
observed the variance of tubercules and
obesity in relationship to stream size in the
Ohio River system (1918: 590) and in the
Green River (1926: 182).

Though Ortmann recognized clinal vari-
ation in torulosa, and Ball carefully docu-
mented it, some taxa ( applied to ecopheno-
typic varients of torulosa) continue to be
promulgated in the literature as if they re-
ferred to subspecies. E. torulosa, like many
other unionid species when found in large
lakes, show some ecophenotypic variation.
Often these forms have been named; in this
case, Dysnomia sulcata delicata Simpson.

Ortmann (1926: 182) mentioned that
\\ alker had specimens of torulosa from
the Cumberland River. "According to the
labels, they have gone through the hands of
W'c'therby and Marsh, and probably were
collected by Dr. Lindsey in 1877." Addi-
tional specimens from this lot in the Mu-
seum of Zoology, University of Michigan
are now in the Museum of Comparative Zo-
ology 236769. Since this species appears to

have been relatively abundant Wiicie lo-
cated, and as it was not found daring the
extensive Cumberland River coliechng re-
ported by Wilson and Clark (1914), this
record is regarded as spurious.

Range. Tennessee River system, Ten-
nessee and Alabama; Ohio River system:
from Illinois to Pennsylvania, including the
Wabash, Green, Licking and Kentucky
River drainages; St. Lawrence River sys-
tem: Lakes Michigan, Huron, and Erie.

Abundance. Still occasionally collected
in commercial operations on the lower Ohio
River ( Kentucky-Ilhnois ) (Parmalee 1967:
62 ) and from the Nolichucky River near its
mouth in western Tennessee. Persists in
smaller streams in the Ohio and lower
Great Lakes systems [Stansbery as D. t.
rangiana] and in the Clinch River, Tennes-
see [Stansbery as D. t. guhernaculum].
Gone throughout the rest of its previous
range (Stansbery, 1970; 20). Epioblasma
torulosa torulosa is listed as "extirpated" by
Stansbery (1976: 43, 51).

Specimens Examined

Tennessee River System

Powell River Drainage. Virginia: Pow-
ell River, Jonesville, Lee Co. (MCZ);
Powell River, Shawanee, Clairborne Co.

(CM).

Clinch River Drainage. Virginia:
Clinch River, Dungannon (CM); Clinch
River, Hill Station, 5.5 mi. below Fort
Blackmore (MCZ); Clinch River, CHnch-
port (MCZ); all Scott Co. Tennessee:
Clinch River, Kyles Ford, Hancock Co.
(MZUM); Clinch River, Union Co.; Clinch
River, Clinton, Anderson Co. (both MCZ).

Holston River Drainage. Virginia:
North Fork, Holston Bridge, Scott Co.
(CM). Tennessee: Holston River, McBee
Ford, Hodges, Jefferson Co.; mouth of
Holston River, Austins Grist Mill, Knox Co.
{both MCZ).

French Broad River Drainage. Tennes-
see: Nolichucky River, 3.5 mi. SE War-
rensburg, Greene Co. (MCZ); Nolichucky
River, Chunns Shoals, Hamblen Co. (CM).

264 Bulletin Museum of Cotnparative Zoology, Vol. 148, No. 6

Tennessee River Drainage. Tentxessee:
Tennessee River near Knoxville, Knox Co.
(MCZ); Tennessee River, Chattanooga,
Hamilton Co. (CM).

Paint Rock River Drainage. Alabama:
Paint Rock River, Poplar Bluff Ridge, Mad-
i'onCo. (MCZ).

Tennessee River Drainage. Ala])ama:
Tennessee River, 6 mi. E Decatur Morgan
Co. (MZUM).

Elk River Drainage. Aluhanui: Elk
River, Fayetteville, Lincoln Co. (MZUM).

Tennessee River Drainage. Alahama:
Tennessee River, Muscle Shoals, Colbert
and Lauderdale Cos. ( MCZ, CM ) ; Tennes-
see Rixer, Florence, Lauderdale Co. ( MCZ,
USNM ) : Tennessee River, Tuscumbia,
Colbert Co. (USNM); Pickwick Basin,
mounds between Barton, Colbert Co. and
Waterloo, Lauderdale Co. ( Morrison,
USNM).

Ohio River System

Ohio River Drainage. Illinois: Ohio
Ri\'er, Metropolis, Nhissac Co. (Parmalee,
1967).

Wabash River Drainage. Indiana: Eel
River, N Manchester, Montgomery Co.
(MCZ); Blue River, Morristown (MZUM);
Conn Creek, Waldron; East Fork Flat Rock
Creek, [town of] Flat Rock Creek (both
MCZ); all Shelby Co. [Mississinewa
River], Albany, Delaware Co. (USNM);
White River, Indianapolis, Marion Co.
(MZUM); White River, Rockford, Jackson
Co. (CM). Wabash River, Delphi; Wild
Cat Creek; both Carroll Co. {])oth MCZ).
Wabash River, Lafayette, Tippecanoe Co.
(MCZ). Wabash River, New Harmony,
Posey Co. (USNM).

Green River Drainage. Kentucky:
Green River, 8 mi. S Campbellsville, Tay-
lor Co. (MCZ); Green River, Greensburg,
Green Co. (MZUM); Mamouth Cave, Ed-
mondson Co. (MCZ); Drakes Creek, 1 mi.
SE Mt. Victor (MZUM); Barren River,
Bowling Green (MCZ); both Warren Co.

Kentucky River Drainage. Kentucky:
Kentucky River ( Rafinesque and Dan-
glade).

Licking River Drainage. Kentucky:

Licking River (MCZ).

Ohio River Drainage. Ohio: Ohio
River, Cincinnati, Hamilton Co. (MCZ).

Little Miami River Drainage. Ohio:
Little Miami River (MCZ).

Scioto River Drainage. Ohio: Sci-
oto River, Columbus, Franklin Co.; Big
Darby Creek, 4 mi. S Orient {both MCZ);
Scioto River, Circleville (USNM); both
Pickaway Co.; Scioto River, Chillicothe,
Ross Co. (USNM).

Muskingum River Drainage. Ohio:
Tuscarawas River, New Philadelphia, Tus-
carwas Co. ( MCZ ) .

Ohio River Drainage. Ohio: Ohio
River, Marietta, Washington Co. (MZUM).

Big Beaver River Drainage. Ohio:
[Little Mahoning River], Newton Falls,
Trumbull Co. (MZUM); Mahoning River,
near Garrettsville, Portage Co. (MCZ);
Mahoning River, near Poland, Mahoning
Co. (Lea). Pennsylvania: Shenango River,
Pulaski and Harbor l^ridge; both Lawrence
Co. {both CM).

Allegheny River Drainage. Pennsyl-
vania: Allegheny River, Warren, Warren
Co. (MCZ); Allegheny River, Hickory and
Tionesta, both Forest Co.; Allegheny River,
Walnut Bend, Venango Co.; French Creek,
Meadville and Cochranton, both Crawford
Co.; French Creek, Utica, Venango Co.;
Allegheny River, Templeton, Johnetta,
Godfrey, and Aladdin; all Armstrong Co.
(all CM).

Monongahela River Drainage. West
Virginia: West Fork Lynch Mines, Harri-
son Co. (CM).

St. Lawrence River System

Great Lakes Drainage. (Lake Michi-
gan) Michigan: Grand River, Grand Rap-
ids, Kent Co. (MCZ). (Lake Huron)
Michigan: Black River, 1.5 mi. W Ama-
dore, Salinac Co. (MZUM). (Lake Erie)

Plagiola from Middle North America • Jc'.-'.iOTi

265

A//c7?/f,'fln: Clinton River, Pontiac, Oakland
Co. (MZUM). Ontario: Syndenham River,
1.8 mi. NE Shetland, Lambton Co.
(Clarke); Detroit River [=Lake Erie], Am-
herstburg (USNM); Detroit River, Bois
Blanc Isle (MZUM); both Essex Co. Mich-
i<l,an: Huron Ri\'er, Huron Park, Wayne
Co.; River Raisin, Petersburg, Monroe Co.
{l)oth MZUM). Indiana: St. Marys River
(MZUM).

Plagiola {Torulosa) sampsoni (Lea)
Plate 12, figures 6, 7
Distribution: Plate 6, figure B

Unio sampsonii Lea 1861, Proc. Acad. Nat. Sci.
Phila. 13: 392 (Wabash River, New Harmony
[Posev Co.], Indiana); 1862, Jour. Acad. Nat.
Sci. Phila. (2)5: 192, pi. 25, fig. 261, figured
holotype UNSM 84802; 1863, Obs. Unio 9: 14.

TninciUa (Pilea) sampsoin (Lea). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 523; 1914, Cat. Na-
iades 1 : 27.

Tniucilla sampsoni (Lea). Ball, 1922, Ecology 3:
117.

Dijsiiomia {Torulosa) sampsoni (Lea). Frierson,
1927, Check list N American naiades, p. 95.
Haas, 1S69, Das Tierreich, pt. 88, p. 486.

Dijsnomia sampsoni (Lea). Goodrich and van der
Schalie, 1944, Amer. Mid. Nat. 32: 314. Parma-
lee, 1967, Illinois State Mus., Popular Sci. Ser.
8: 92, pi. 30 E. Stansbery, 1970, Malacologia
10: 20, pi. 2, fig. 6; 1971, Symposium of rare
and endangered moll. U. S., p. 18b, fig. 18.

Description. Shell of medium size, sel-
dom reaching over 50 mm in length. Out-
line subquadrate or trapezoid. Valves in-
equilateral, inflated, solid. Anterior end
regularly rounded, posterior end of the male
somewhat pointed, rather broadly rounded
in females. Ventral margin straight or
slightly curved. Dorsal margin usually suf-
ficiently straight to form an angle with the
obliquely descending posterior slope, but
sometimes rounded and join'ng the pos-
terior margin imperceptibly. Hinge liga-
ment short. Posterior ridge low, narrowly
rounded, separated from a somewhat
broader medial ridge by a distinct furrow
that ends in an emargination between the
somewhat produced ridges in most males,

though in some males the furrow ends in
a straight line; in females it is broadly
rounded. The ridges are occasionally slight-
ly knobbed, but the furrow is always
smooth. The marsupial swelling in tfie fe-
male is infrequently marked by a miinher
of small radial furrows. The rounded mar-
supial swelling extends from the middle of
the base to the upper part of the posterior
end. Umbos high and full, turned over a
small wide lunule, located toward the an-
terior third of the shell, their sculpture
feeble and corrugated. Surface of the shell
rather smooth, usually with a concentric
constriction at the rest marks. Periostracum
smooth, usually shiny, though sometimes
s Iky, yellowish green or straw colored with

numerous green ravs.

Left Xcdve with two triangular pseudo-
cardinal teeth; slight interdentum. Two
moderately long, almost straight, lateral
teeth. Right valve with one large pseudo-
cardinal tooth with a smaller tooth before
and after it. One lateral tooth, sometimes
with a vestigial tooth below. Umbonal cavi-
ty shallow. Anterior and posterior muscle
scars and pallial line well impressed. Nacre
bluish white to salmon.

Usually female shells do not differ great-
ly in outline from the male. The female
marsupial swelling is usually just a little
produced, not differing in color or texture
from the rest of the shell.

Length Height Width

Lower Wabash River, In-
diana. Male. (USNM).

As above. Female.

Hahitat. Lixes in sand and gravel bars,
never in mud. ( Lea ) .

Remarks. Pla<i,iola .samp.soni (Lea) most
closely resembles P. torulosa (Rafinesque),
but it differs from the latter in that it does
not reach such large size, it is almost always
devoid of knobs, and both the male and fe-
male shells are subquadrate or trapezoid in
outline (a characteristic that does not vary

266 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

imicli because of sexual differences). The
outline of the male torulosa is ovate and
that of the female obovate; the marsupial
swelling of the female is darker than the
rest of the shell, whereas in sampsoni it is
the same color as the rest of the shell. P.
mmpsoni also resembles P. propinqua in
that it is generally without knobs. How-
e\'er, male shells of the latter are subtri-
angular in outline while those of the female
are subelliptical. The periostracum of
sampsoni is generally shiny while that of
propinqua is almost always subshiny.

Call (1900: 476) included sampsoni un-
der the synonymy of Unio perplexus [ =
torulosa]. Goodrich and van der Schalie
(1944: 314) suggested that sampsoni might
be a large river form or variant of Dys-
nomia perplexa, though they admitted
there were characters that might entitle it
to specific rank. Stansbery (1970: 20, pi.
2, fig. 6) thought sampsoni might be sim-
ply a variant of the subspecies he recog-
nized as Dysnomia torulosa rangiana (Lea).
As pointed out above, both the male and
female shell of sampsoni have sufficient
characters to separate it from the other
Plagiola.

Range. Tennessee River system, Tennes-
see. Ohio River system: lower Wabash
River drainage, Indiana; Ohio River drain-
age to Cincinnati, Hamilton Co., Ohio.

Abundance. This species was not men-
tioned by Ortmann ( 191S, 1925) or Morri-
son ( 1942 ) in their studies of the unionids
of the Tennessee River system. Ball (1922:
117) had only three specimens without lo-
cality data, which could not be presently
located in the Carnegie Museum. It is pre-
sumed to be known from the Tennessee
River system, from a male and female
specimen in the Museum of Comparative
Zoology merely labeled, "Tennessee."

Formerly found in fair numbers in the
lower Wabash River, Indiana. Not located
there by Meyer (1974) or Clark (1976).
Considered extinct by Stansbery (1970:
20).

Specimens Examined

Tennessee River System

Tennessee River Drainage. Tennessee:
Tennessee (MCZ. LocaHty as well as the
collector. Steward, written on the male and
female shells ) .

Ohio River System

Wabash River Drainage. Indiana:
White River (MZUM). Wabash River,
New Harmony (MZUM, USNM, ANSP);
Grand Chain (USNM); both Posey Co.
Illinois. Wabash River, 'Little Chains,'
[archaeological site], 10 mi. above con-
fluence with the Ohio River, White Co.
(Parmalee).

Ohio River Drainage. Ohio: Ohio
River (ANSP); Ohio River, Cincinnati,
Hamilton Co. (MCZ).

Plagiola (Torulosa) propinqua (Lea)
Plate 12, figures 8, 9
Distribution: Plate 6, figure B

Unio propinquus Lea, 1857, Proc. Acad. Nat. Sci.
Phila. 9: 83 ([Tennessee River], Florence [Lau-
derdale Co.]; [Tennessee River] Tuscumbia
[Colbert Co.]; both Alabama); 186L Jour. Acad.
Nat. Sci. Phila. (2) .5: 63, pi. 5, fig. 212, figured
holotype USNM 84332 from the former locality;
1862, Obs. Unio 8: 67.

TninciUa (Pilea) propinqua (Lea). Simpson,
1900, Proc. U. S. Natl. Mus. 22: 523; 1914, Cat.
Naiades 1: 27.

TninciUa propinqua (Lea). Ortmann, 1918, Proc.
Amer. Pbilos. Soc. 57: 589. Ball, 1922, Ecology
3: 113.

Dysnomia (Pilea) torulosa propinqua (Lea). Ort-
mann, 1925, Amer. Mid. Nat. 9: 363.

Dysnomia (Torulosa) propinqua (Lea). Frierson,
1927, Cbcck list N American naiades, p. 95.
Haas, 1969, Das Tierreich, pt. 88, p. 487.

Dysnomia propinqua (Lea). Morrison, 1942, Bur.
Amer. Etbnology, Bull. no. 129, p. 365. Stans-
bery, 1970, Maiacologia 10: 20, pi. 2, fig. 5;
1971, Svmposium of rare and endangered moll.
U. S., p.' 18b, fig. 17.

Description. Shell of medirmi size, sel-
dom reaching over 60 mm in length. Out-
line subtriangular, subelliptical or subquad-
rate. Valves very inequilateral, inflated,

Plagiola from Middle North America ' Joh.r.sD^ 267

solid. Anterior end regularly rounded, pos-
terior end of male more broadly rounded,
sometimes somewhat pointed; posterior
more broadly and evenly rounded in fe-
males. \^entral margin curved. Dorsal mar-
gin sometimes sufficiently straight to form
an angle with the obliquely descending
posterior margin, but often rounded and
joining the posterior margin imperceptibly.
Hinge ligament short. Posterior ridge low,
narrowly rounded, median ridge full or
rounded, sometimes the ridges are slightly
knobbed. Dorsal slope usually rounded.
The radial furrow that separates the ridges
varies in both depth and width, but it is
usually rather deep, especially in the male.
Umbos full and high, somewhat turned over
a small wide lunule, located in the anterior
quarter of the shell, their sculpture is feebly
corrugated. Surface of the shell with many
distinct growth rests. Periostracum vary-
ing from straw-colored or tawny to yellow
green with feeble green rays, subshiny or
somewhat silky.

Left valve with two chunky triangular
pseudocardinal teeth; slight interdentum.
Two short, slightly curved lateral teeth.
Right valve with one large triangular pseu-
docardinal tooth usually with a smaller
tooth before and behind it. One lateral
tooth, sometimes with a vestigial tooth be-
low. Umbonal cavities shallow. Anterior
and posterior adductor muscle scars, and
pallial line, well impressed. Nacre bluish
white, rarely pinkish.

Male shells subtriangular, with the radial
furrow usually deep, ending in a narrow
sinus below.

Female shells subelliptical or subquad-
rate, the marsupial swelling rounded and
moderately produced, the radial furrow be-
coming obscure posteriorly. Marsupial
swelling sometimes darker than the rest of
the shell.

Remarks. Plagiola propinqua (Lea)
bears some resemblance to P. torulosa, and
though the former may be slightly knobbed,
it is generally smooth. The male of propin-
qua is subtriangular in outline, while that of

torulosa is generally irregularly o /ate. The
posterior ridge of the former is more
obliquely angled, and the radial fuiTOw is
narrower and deeper than in torulosa. I'be
female of propinqua is subelliptical in out-
line, while the torulosa female is produced
posteriorly and obovate. The radial furrow
of propinqua is narrow and deep except in
the post-basal region where there is some
marsupial swelling. The posterior end of
the female torulosa is without a radial fur-
row, but is instead a large flattened mar-
supial swelling that extends from the mid-
dle of the ventral margin to the upper part
of the posterior margin. While the mar-
supial area of propinqua is usually the same
color as the rest of the shell, that of torulosa
is often a darker green.

Simpson (1914: 24) correctly recognized
propinqua as a distinct species, but Ort-
mann (1925: 363) in interpreting the data
of Ball (1922: 115) regarded propinqua as
merely a torulosa with the tubercules very
poorly developed or absent. Morrison
( 1942: 365) examined thousands of torulosa
and propinqua specimens from Indian
mounds of the Pickwick Landing Basin,
Colbert and Lauderdale counties, Alabama
and found no integrades between them.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem, Tennessee; Ohio River system: from
the lower Wabash River drainage, Indiana
to the Ohio River, Cincinnati, Hamilton
County, Ohio.

Abundance. This species was not found
by Wilson and Clark (1914) in the Cum-
berland River svsteni. Considered extinct
by Stansbery (1976: 43, 51).

Specimens Examined
Tennessee River System

Clinch River Drainage. Tennessee:
Clinch River. Clinton and Edgemoor, both
Anderson Co. {both CM).

Hoist on River Drainage. Tennessee:
mouth of Holston Ri\'er, Austins Grist Mill,
Knox Co. (MCZ; MZUM).

268

Bulletin ^[ll.scum of Comparative Zoology, Vol. 148, No. 6

Tennessee River Drainage. Tennessee:
Tennessee River, Knoxville, Knox Co.
(MCZ); Alabama: Tennessee River, Mus-
cle Shoals, Colbert and Lauderdale Cos.
(CM); Tennessee River, Florence, Lauder-
dale Co. (MCZ, MZUM); Tennessee
River, Tuscumbia, Colbert Co. (MCZ);
Pickwick Basin, mounds between Barton,
Colbert Co. and \\'aterloo, Lauderdale Co.
(Morrison, USNM).

Cumberland River System

Cumberland River Drainage. Tennes-
see: Cumberland River (Simpson, 1914:
27; MZUM; MCZ). [Cumberland River],
Nashville, [Davidson Co.] (MCZ, data
written on the shells).

Ohio River System

Wabash River Drainage. Indiana:
White River (MZUM); Wabash River,
New Harmony, Posey Co. (MCZ; MZUM).

Ohio River Drainage. Ohio: Ohio
River, Cincinnati, Hamilton Co. (MCZ).

Plagiola (Toruiosa) biemarginata
Plate 13, figures 1, 2
Distribution: Plate 9, figure A

(Lea)

Viiio biemarginatus Lea 1857, Proc. Acad. Nat.
Sci. Phila. 9: 83 ( [Tennessee River], Florence
[Lauderdale Co.], Alabama); 1866, Jour. Acad.
Nat. Sci. Phila. (2) 6: 47, pi. 16, fig. 45, figured
holotype USNM 84608; 1867, Obs. Unio 11: 51.

Tnincilla (Pilea) biemarginata (Lea). Simpson,
1900, Proc. U. S. Natl. Mus. 22: 524; 1914, Cat.
Naiades 1: 28.

Dysnomia (Pilea) biemarginata (Lea). Ortmann,
1925, Amer. Midland Nat. 9: 361.

Dysnomia (Toruiosa) biemarginatus (Lea). Frier-
son, 1927, Check list N American naiades, p.
95. Haas, 1969, Das Tierreich, pt. 88, p. 487.

Dysnomia biemarginata (Lea). Morrison, 1942,
Bur. Amer. Ethnology, Bull. no. 129: 364. Stans-
bery, 1970, Malaco'logia 10: 20; 1971, Sym-
posium of rare and endangered moll. U. S., p.
18b, figs. 13, 14.

Description. Shell usually small, seldom
reaching more than 50 mm in length. Out-
line subtriangular, subrhomboid, or irregu-

larly obovate. Valves not much inflated,
solid, inequilateral. Anterior end regularly
rounded, posterior end of male less broadly
rounded, subtruncated; more broadly and
more evenly rounded in females. Ventral
margin of male curved, of female almost
straight before the marsupial swelling. Dor-
sal margin of male long, forming a sharp
angle with the obliquely descending pos-
terior margin; of female short forming a less
sharp angle with the obliquely descending
posterior margin. Hinge ligament rather
long. Posterior ridge of male with a sharp,
biangulate, posterior ridge ending in a bi-
angulation below, median ridge full and
rounded, the radial depression between the
ridges wide and shallow. Dorsal slope con-
cave. Posterior ridge of the female faintly
biangulate, the radial depression between it
and the medial ridge shallow and fading out
on the marsupial swelHng. Umbos moder-
ately full and inflated, located sHghtly an-
terior of the middle, their sculpture not ob-
served. Surface of the shell with strong and
irregular growth rests. Periostracum yel-
lowish green, with numerous green rays of
varying width over the entire surface.

Left \'al\'e with two chunky triangular
pseudocardinal teeth; slight interdentum.
Two long, almost straight, lateral teeth.
Right valve with one large triangular
pseudocardinal tooth, usually with a much
smaller tooth before and behind it. One
lateral tooth. LTmbonal cavities shallow.
Anterior and posterior adductor muscle
scars and pallial line well impressed. Shell
thinner posteriorly in both sexes. Nacre
bluish white to creamy.

Male shells are subtriangular, or sub-
rhomboid, with a sharply biangulate pos-
terior ridge that ends posteriorly in a bi-
angulation. There is a wide shallow radial
depression before a full rounded medial
ridge.

Female shells are obovate, having a large
flattened, rounded marsupial swelling ex-
tending from the middle of the base to the
upper part of the posterior end. The radial

Plagiola from Middle North America • Jc'r/nscn 269

depression between the faintly biangulate
posterior ridge and the medial ridge is shal-
low and fades out on the marsupial swelling
which is darker than the rest of the shell.

Length Height Width
mm mm mm

Tennessee River, Muscle
Shoals, Colbert and Lau-
derdale Cos., Alabama.
Male.

As above. Female.

Remarks. Plagiola biemarginata (Lea)
bears a resemblance to both P. perplexa and
propinqua. It is smaller than both of them,
and is not knobbed as perplexa often is. It
may be distinguished easily from propin-
qua, which has a much higher shell in re-
lation to length. W'hile close to the same
proportions as perplexa, biemargitmta is
easily separated from it by the decidedly
more acutely biangulate posterior ridge,
which ends in a sharper biangulation at the
posterior end in males, and has a concave
dorsal slope. The wide green rays on both
sexes of biemarginata distinguish it from
perplexa and propinqua.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem. Big South Fork, Kentucky.

Abundance. Considered extinct by Stans-
bery (1970: 20; 1976: 43, 50).

Specimens Examined
Tennessee River System

Clinch River Drainage. Tennessee:
Clinch River, "The Rounds," Hancock Co.
(MCZ).

Holston River Drainage. Tennessee:
Holston River, Knox Co. (MCZ).

Sequatchie River Drainage. Tennessee:
Sequatchie River (MZUM).

Paint Rock River Drainage. Alabama:
Paint Rock River, Paint Rock, Jackson Co.
(MZUM).

Flint River Drainage. Alabama: Flint
River (MZUM).

Elk River Drainage. Tennessee. Elk
River, Fayetteville, Lincoln Co. (MZUM).

Tennessee River Drainage. Alabama:
Tennessee River, Muscle Shoals, Colbert
and Lauderdale Cos.; Tennessee River,
Florence, Lauderdale Co.; Tennessee River,
Tuscumbia, Colbert Co.; {all xMCZ); Pick-
wick Basin, mounds between Barton, Col-
bert Co. and Waterloo, Lauderdale Co.
(Morrison, USNM).

Cumberland River System

Cumberland River Drainage. Ken-
tuchj: Big South Fork, Burnside, Pulaski
Co. (MZUM).

Plagiola (Torulosa) capsaeformis (Lea)
Plate 13, figures 3, 4
Distribution: Plate 7, figure B

Unio capsaeformis Lea 1834, Trans. Amer. Philos.

Soc. 6: 31, pi. 2, fig. 4 (Cumberland River

[Tennessee], figured holotype MCZ 178570);

1834, Obs. Unio 1 : 143.
TninciUa (Pilea) capsaeformis (Lea). Simpson,

1900, Proc. U. S. Natl. Mus. 22: 524; 1914, Cat.

Naiades 1 : 29.
TrunciUa capsaeformis (Lea). Ortmann, 1912,

Ann. Carnegie Mus. 8; 359 [anatom\- mistakenly

described under florentina]; 1913, Proc. Amer.

Philos. Soc. 52: 311. Goodrich, 1913, Nautilus

27: 95.
Dijsnomia (Pilea) capsaeformis (Lea). Ortmann,

1924, Amer. Mid. Nat. 9: 38; 1925, Amer. Mid.

Nat. 9: ;362.
Dijsnomia (Capsaeformis) capsaeformis (Lea).

Frierson, 1927, Check list N American naiades,

p. 95. Haas, 1969, Das Tieneich, pt. 88, p. 487.
Dijsnomia capsaeformis (Lea). Wilson and Clark,

1914, U. S. Bur. Fisheries, Doc. 781, p. 46.

Morrison, 1942, Bur. Amer. Ethnology, Bull.

no. 129, p. 364. Neel and Allen, 1964, Mala-

cologia 1: 448. Van der Schalie, 1973, Sterki-

ana, no. 52, pp. 46, 48, 50, 51.
Epiobtasma capsaeformis (Lea). Stansbery, 1972,

Amer. Mai. Union, Bull, for 1972, p. 22.

Description. Shell of medium size, reach-
ing about 70 mm in length. Outline ellip-
tical or irregularly obovate. Valves some-
what inequilateral, subinflated, subsoHd.
Anterior end regularly rounded, posterior
end of male slightly produced; more broad-

270 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

ly rounded in females. Ventral margin
slighth-, but uniformly, curved in the male;
almost straight in females to the sulcus,
behind which the marsupial swelling ex-
tends well below the base. Dorsal margin
straight, forming an angle with the
obliquely descending posterior margin.
Hinge ligament short. Posterior ridge of
the male, double, but faint, ending in a
slight biangulation posteriorly; ridge
scarcely visible in females. There is some-
times a faint radial depression in front of
the lower posterior ridge of the male. The
considerable marsupial swelling of the fe-
male is usually marked by a sulcus before
and behind and is sometimes finely toothed
on the margin. Umbos quite full and ele-
vated, located slightly anterior of the mid-
dle in the male, and in the anterior third of
the shell in the female; umbonal sculpture
not observed. Surface of the shell with un-
even growth lines. Periostracum subshiny,
yellowish green with numerous fine green
rays over the entire surface, marsupial area
of the female usually dark green, sometimes
blackish.

Left valve with two, chunky, triangular
pseudocardinal teeth; slight interdentum.
Two short, slightly curved lateral teeth.
Right valve with one large triangular
pseudocardinal tooth, usually with a smaller
tooth before it that is parallel to the hinge
line. One lateral tooth, sometimes with a
vestigial tooth below. Umbonal cavities
shallow. Anterior adductor muscle scars
well impressed, posterior ones shallow. Pal-
lial line distinct anteriorly. Nacre bluish
white. Shell heavier anteriorly, posterior
end of females especially thin and irri-
descent.

Male shells irregularly elliptical, with a
double, but faint, posterior ridge, which
ends in a slight biangulation, near the me-
dian.

Female shells irregularly obovate with
a thin, slightly inflated, marsupial swelHng,
which may be considerably produced and
extended well below the base, often marked

by two distinct sulci, sometimes toothed on
the margin.

Length Height Width
mm mm mm

Duck River, Milltown,
Marshall Co., Tennessee.
Male.

As above. Female.

Remarks. Plagiola capsaefonnis (Lea)
bears some resemblance both to P. perplexa
and florentina. Only the female resembles
perplexa, but that of capsaeformis is always
smaller, thinner and without knobs. Both
the male and female of capsaeformis resem-
ble florentina, but the male of the former
is longer, lower and less swollen than that
of florentina. In the capsaeformis female
the marsupial swelling is darker than the
rest of the shell, while in florentim the
periostracum (in both sexes) is a uniform
honey yellow or yellowish brown. The
green rays of the latter are more evenly dis-
tributed.

Range. Tennessee River system, Vir-
ginia, Tennessee and Alabama; Cumber-
land River system, Kentucky and Tennes-
see.

Abundance. The number of specimens
seen in collections indicates this species
must have been abundant, especially in the
Tennessee River system.

Specimens Examined
Tennessee River System

Powell River Drainage. Virginia: Pow-
ell River, 2.5 mi. S Jonesville, Lee Co. Ten-
nessee: Powell River, Shawnee (CM);
Powell River, 8-10 mi. N Tazewell (MCZ);
both Claiborne Co.; Powell River, Greens
Ford, 2 mi. NW Long Hollow, Union Co.
(MCZ).

Clinch River Drainage. Virginia: Clinch
River, Cedar Bluff, Tazewell Co. (CM);
Clinch River, Cleveland, Russell Co.
(MCZ); CUnch River, Dungannon, Clinch-
port, and below Speers Ferry Bridge, all
Scott Co. (all MCZ); Clinch River, 1.5 mi.

Plagiola from Middle North America • Johnson

S Dona, Lee Co. (MCZ). Tennessee:
Clinch River, Kyles Ford, and "The
Rounds," ])Oth Hancock Co. Clinch River,
4 mi. NW Thorn Hill, Grainger Co.;
Clinch River, Clinton, Anderson Co. (all
MCZ).

Holston River

Drainage.

Virainia:

North Fork, Mendota, Washington Co.
(CM); North Fork, Hilton, Scott Co.
(MCZ). Tennessee. North Fork, Clouds
Ford, 2 mi. W Morrison (MCZ); South
Fork, Pactolus (CM); both Sullivan Co.;
mouth of Holston River, Austins Grist Mill,
Knox Co. (MCZ).

French Broad River Drainage. North
Carolina: French Rroad River, Asheville,
Buncombe Co. (MZUM). Tennessee:
Nolichucky River, 3.5 mi. SE Warrensburg,
Green Co.

Tennessee River Drainage. Tennessee:
Tennessee River, near Knoxville, Knox Co.
(both MCZ); Little Tennessee River, Coy-
tee, Loundon Co. (MZUM).

Paint Rock River Drainage. Alabama:
Paint Rock River, Princeton, Holly Tree,
and Trenton (all MCZ); between New
Hope and Paint Rock (CM); all Jackson
Co.

Flint River Drainage. Alabama: Flint
River, Gurley, Madison Co. (MCZ).

Limestone Creek Drainage. Alabama:
Limestone Creek, Mooresville, Limestone
Co. (MZUM).

Elk River Drainage. Tennessee: Elk
River, Winchester, Franklin Co. (MCZ);
Richland Creek, Wales, Giles Co. (CM).
Alabama: Elk River, Fayetteville, Lincoln
Co. (MCZ).

Tennessee River Drainage. Alabama:
Shoals Creek, Lauderdale Co.; Tennessee
River, Muscle Shoals, Colbert and Lauder-
dale Cos. (both MCZ); Pickwick Basin,
mounds between Barton, Colbert Co. and
Waterloo, Lauderdale Co. (Morrison,
USNM).

Bear Creek Drainage. Alabama: Bear
Creek, Burleson, Franklin Co. (CM).

Duck River Drainage. Tennessee: Duck
River, Shelbyville, Bedford Co. (MZUM);

Duck River, Wilhoite; Duck River, below
Lillard Mill, Miltown; Duck River, Har-
dinsons Mill, 12 mi. NW Lewisburg; all
Marshall Co. (all MCZ); Duck River, Left-
wich (CM) and Columbia (MCZ), both
Maury Co.

Cumberland River System

Cumberland River Drainage. Ken-
tucky: Cumberland River, below Cumber-
land Falls, Whitely Co.; Rockcastle River,
Mt. Victory; (both MZUM); Big South
Fork, Parkers Lake Station (Wilson and
Clark) and above Burnside (MZUM); all
Pulaski Co.; Cumberland River, Horseshoe
Bottom (MZUM); Beaver Creek, E Ro-
wena Ferry (MCZ); both Russell Co. Cum-
berland River, Neeleys Ferry, Cumberland
Co. (MZUM).

Obey River Drainage. Tennessee: Obey
River, Pryor Bend, Pickett Co. (MCZ).

Caney Fork Drainage. Tennessee: Ca-
ney Fork [Smith Co.], (Wilson and Clark).

Harpeth River Drainage. Tennessee:
Haipeth River, 10 mi. W Franklin, William-
son Co.; Harpeth River, Davidson Co.
(both MCZ).

Plagiola (Torulosa) florentina (Lea)
Plate 13, figures 5-14
Distribution: Plate 2

Unio florentiniis Lea 1857, Proc. Acad. Nat. Sci.
Phila. 9: 83 ([Tennessee River], Florence,
[Lauderdale Co.], Alabama; Cumberland River,
Tennessee); 186L Jour. Acad. Nat. Sci. Phila.
(2) 5: 64, pi. 5, fig. 213, figured holotype
USNM 84948 from the former locality; 1862,
Obs. Unio 8: 68.

Unio saccatus Kiister 1862, Conch. Cabinet ( 2 )
9, pt. 2, p. 263, pi. 89, fig. 2 (Tennessee, figured
type, location unknown ) .

Unio saccuhis Reeve 1864, Conch. Iconica 16,
Unio, pi. 15, species 67 ( North America, figured
type, British Mus. Nat. Hist. [lost]). Anthony,
1865, Amer. Jour. Conch. 1: 157, pi. 12, fig. 3
(Tennessee, figured holotype MCZ 161898).
Anthony sent H. Cuming a specimen of his
species before publication, which Reeve de-
scribed before Anthon>-.

Tnincilla walkeri Wilson and Clark 1914, U. S.
Bur. Fisheries. Doc. no. 781, p. 46, pi. 1, fig. 1

272 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

(East Fork of Stones Ri\er, near Walterville
[rzWalterliill, Rutherford Co.], Tennessee, since
the two figured s\ntypes could not be located
in the USNM, a Icctotypc is here selected from
a series of syntypes in the Museum of Zoology,
University of Michigan 90729, pi. 13, fig. 9).
Ortmann, 1918, Proc. Amcr. Philos. Soc. 57:
592.

Truncilhi curtisii Frierson and Uttcrback 1916,
Amer. Mid. Nat. 4: 453 (190), pi. 6, fig. 14a-d,
pi. 28, fig. 109 A-D (White River, Hollister,
[Taney Co.], Missouri, the two figured syn-
types are lost, a lectotype is here selected from
a series collected by Frierson and Utterback in
the White Ri\er, Forsyth, Taney Co., Missouri,
Museum of Zoology, University of Michigan
90748, pi. 13, fig. 10).

Tniurilla (Pilea) florcntina (Lea). Simpson,
1900, Proc. U. S. Natl. Mus. 22: 524 [partim];
1914, Cat. Naiades 1: 30 [partim]. Wilson and
Clark, 1914, U. S. Bur. Fisheries, Doc. 781, p.
46.

Dysnomia (Pilea) florcntina (Lea). Ortmann,
1925, Amer. Mid. Nat. 9: 362.

Dijf/numia {Capsaeformis) florcntina (Lea). 1927,
Check Hst N American naiades, p. 95. Haas,
1969, Das Tierreich, pt. 88, p. 488.

Description. Shell of medium size, sel-
dom reaching over 60 mm in length. Out-
line elliptical or irregularly obovate. Valves
somewhat inequilateral, subinflated, sub-
solid. Anterior end regularly rounded,
posterior end of male slightly produced;
posterior more broadly rounded in females.
Ventral margin slightly but uniformly
curved in the male; almost straight in fe-
males to the sulcus, behind which the mar-
supial swelling extends well below the
base. Dorsal margin straight, forming an
angle with the obli(iuely descending pos-
terior margin. Hinge ligament short. Pos-
terior ridge of the male, double, but faint,
ending in a slight biangulation posteriorly;
ridge scarcely visible in females. There is
a wide shallow radial depression in front
of the full, lower posterior ridge in the
male. The considerable marsupial swelling
of the female is usually marked by a sul-
cus before and behind and is sometimes
finely toothed on the margin.

Umbos quite full and elevated, located
slightly anterior of the middle in the male,
and in the anterior third of the shell in the

female, imibonal sculpture not observed.
Surface of the shell with uneven growth
lines. Periostracum subshiny, yellow, honey
yellow, brownish yellow or whitish with
ninnerous green rays more or less uniformly
distributed over the entire surface.

Left valve with two chunky triangular
pseudocardinal teeth; slight interdentum.
Two short slightly curved lateral teeth.
Right xalve with one large triangular
pseudocardinal tooth, usually with a smaller
tooth before it that is parallel to the hinge
line. One lateral tooth, sometimes with a
vestigial tooth below. Umbonal cavities
shallow. Anterior adductor muscle scars
well impressed, posterior ones shallow. Pal-
lial line distinct anteriorly. Nacre bluish
white. Shell heavier anteriorly, posterior
end of females especially thin and irri-
descent.

Male shells irregularly elliptical, with a
double posterior ridge that ends in a slight
bimargination near the median. There is a
wide, shallow, radial depression in front of
the full, lower posterior ridge.

Female shells irregularly obovate with a
thin, slightly inflated marsupial swelling,
which may be considerably produced and
extended well below the base, often marked
by two distinct sulci, toothed on the margin.

Length Hei«ht Width
mm mm mm

Stones River, 7 mi. NE
Murfreesboro, Rutherford
Co., Tennessee. Male.

As above. Female.

Remarks. Phi<i,ioIa florentina (Lea) most
closely resembles P. capsaeformis. The
male of the former is shorter, higher and
more swollen than that of capsaeformis and
has a distinct wide shallow radial depres-
sion in front of the lower posterior ridge.
This depression is very faint or wanting in
capsaeformis. The female florentina has
more strongly developed and more numer-
ous denticulations on the margin of the
marsupial expansion, and is more convex
than capsaeformis. The two species are
most readily separated by the color of the

Plagiola from Middle North America • Johnson 273

surface — florentina is light yellow, honey
colored, yellowish brown, or whitish, and
is rather uniformly rayed over the entire
sinface; copsaefonnis is green or greenish
brown, and while the male is rather uni-
formly rayed, the marsupial area of the fe-
male is a darker green.

Ortmann (1918: 592) noted that walkeri
was a large, compressed florentina, and that
it was probably only an ecophenotypic vari-
ant. He later \l92'4a: 36) concluded from
the measurements of numerous samples that
the obesity or diameter of the shell changed
with the size of the river — the larger rivers
have the more swollen florenthm, and the
smaller rivers have the more compressed
walkeri. In spite of Ortmann's observa-
tions, walkeri has continued to be promul-
gated in the literature as if it had nomen-
clatorial standing.

Truncilla ciuiisi Frierson and Utterback
from the Ozark Plateau was recognized by
Stansbery (1971: 18e) as a subspecies on
the basis that his specimens were nearly
white (pers. comm. ). Utterback (1916:
453 [190]) mentioned that the epidermis of
his specimens was yellowish brown and
finely and obscurely radiate all over. Be-
cause of its geographical isolation, cutiisi
might be recognized as a subspecies as pres-
ently understood, but this does not seem
warranted on morphological differences.

Anatomy. Described by Utterback
(1916: 453 [190]).

Range. Upper White River system, Mis-
souri; Tennessee River system, Virginia,
Tennessee, and Alabama; Cumberland
River system, Kentucky and Tennessee.

Abundance. "Now restricted in the Cum-
berland River system to the lower Stones
and Red Rivers [as D. /. walkeri] and ap-
parently gone from the entire Tennessee
River system save the South Fork of the
Holston in Virginia," (Stansbery 1970: 20).
Collected in small numbers in the Black
River, Missouri by C. B. Stein in 1964.
Listed as "extirpated" by Stansbery (1976:
43,51).

Specimens Examined
White River System

White River Drainage. Missouri: White
River, HolHster, (Utterback); White River,
Forsyth, (MZUM); both Taney Co.

Black River Drainage. Missouri: Black
River, Hendrickson, Butler Co. (MCZ,
OSM).

Tennessee River System

Holston River Drainage. Virginia: Mid-
dle Fork, Chilhowie, Smyth Co. (CM;
Stansbery, OSM); Middle Fork, 3.7 mi. S
Glade Spring (MCZ); South Fork, Barron
(CM); both Washington Co. Tennessee:
South Fork, Emmett, Sullivan Co.; Holston
River, Holston Station, Grainger Co.; Hols-
ton River, Knox Co. {all CM).

Flint River Drainage. Alabama: Flint
River, Maysville and Gurley; Hurricane
Creek, Gurley; all Madison Co. {all CM).

Limestone Creek Drainage. Alabama:
Limestone Creek, Mooresville, Limestone
Co. (MZUM).

Tennessee River Drainage. Alalmnui:
Tennessee River, Muscle Shoals, Colbert
and Lauderdale Cos. (CM); Cypress
Creek, Florence, (MZUM); Tennessee
River, Florence, (MCZ); both Lauderdale
Co. Pickwick Basin, mounds between Bar-
ton, Colbert Co. and Waterloo, Lauderdale
Co. (Morrison, USNM).

Bear Creek Drainage. Alabama: Bear
Creek, Burleston, Franklin Co. (MZUM).

Duck River Drainage. Tennessee: Duck
River, Lillards Mills, Wilhoite, Marshall Co.
(CM); Duck River, Columbia, Maury Co.
(MCZ).

Cumberland River System

Cunil>erlan<l River Drainage. Ken-
tucky: Buck Creek, 2.1 mi. NE Mt. Victory
(MCZ); Cumberland River, Burnside
(CM); both Pulaski Co.; Beaver Creek,
near mouth, (MZUM); Cumberland River,
Indian Creek Bar (Wilson and Clark);

274 Bulletin Museum uf Comparative Zoology, Vol 148, No. 6

Cumberland River, Horseslioe Bottom, 5
mi. S Jamestown (MZUM); all Russell Co.

Obey River Drainage. Tennessee: Obey
River, Duncan Ford, 4 mi. SE Lilydale,
Pickett Co. (MCZ).

Stones River Drainage. Tennessee: East
Fork, Stones River, W'alterhill; Stones
Ri\er, 7 mi. NNE Murfreesboro; both
Rutherford Co. {both MCZ); Stones River,
1.2 mi. W Couchville, Davidson Co.
(MCZ).

Harpetli River Drainage. Tennessee:
Harpeth River, Belle view, Davidson Co.
(CM).

Red River Drainage. Tennessee: Lower
Red River [Montgomery Co.] (Stansbery,
OSM).

Ptagiota (Torutosa) turgiduta (Lea)
Plate 14, figures 1-6
Distribution: Plate 2

Unio turgiduhis Lea 1858, Proc. Acad. Nat. Sci.
Phila. 10: 40 (Cumberland River, Tennessee;
[Tennessee River], Florence, [Lauderdale Co.],
Alabama); 1861, Jour. Acad. Nat. Sci. Phila.
(2) 5: 62, pi. 5, fig. 211, figured holotype
USNM 84946 from the former locality; 1862,
Obs. Unio 8: 66.

Unio mix Kiister 1861, Conch. Cabinet (2) 9, pt.
2, p. 218, pi. 73, fig. 2 (Alabama, figured type,
[location unknown] ) iion Lea 1852.

Unio deviatiis Reeve 1864, Conch. Iconica 16,
Unio, pi. 15, species 61 ( [Tennessee], figured
holotype British Mus. Nat. Hist. 1965210).
Anthony, 1865, Amer. Jour. Conch. 1: 156, pi.
12, fig. 2 (Tennessee, figured holotype MCZ
161895). Anthony sent H. Cuming a si^ecimen
of liis species before publication, which Reeve
described before Anthony.

Truncilla lefevrei Utterback 1916, Amer. Mid. Nat.
4: 455 [192], pi. 6, figs. 13 a-d, pi. 28, figs.
108 A-D (Black River, Williamsville, [Wayne
Co.], Missouri, figured syntypes [lost]).

Truncilla (Pilea) deviata (Reeve). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 524; 1914, Cat. Na-
iades 1 : 32.

Truncilla (Pilea) florcntina (Lea). Simpson,
1900, Proc. U. S. Natl. Mus. 22: .524 [partim];
1914, Cat. Naiades 1: 32 [partim].

Truncilla turgidula (Lea). Ortmann, 1918, Proc.
Amer. Philos. Soc. 57: 590.

Dijsnomia (Pilea) turgidula (Lea). Ortmann and

Walker, 1922, Occ. Papers, Mus. Zool., Univ.
Michigan no. 112, p. 69. Ortmann, 1924, Amer.
Mid. Nat. 9: 34; 1925, Amer. Mid. Nat. 9: 361.

Dijsnoniiu (Capsaeformis) turgidula (Lea). Frier-
son, 1927, Check list N American naiades, p.
95. Haas, 1969, Das Tierreich, pt. 88, p. 490.

Dy.moniia hicmarginata turgidula (Lea). Stans-
bery, 1970, Nhdacologia 10: 20.

Dy.snoniia turgidula (Lea). Stansbery 1971, Sym-
posium of rare and endangered moll. U. S., p.
18b, fig. 19. Van der Schalie, 1973, Sterkiana
no. 52, p. 52.

Dysiwniia (Capsaeformis) lefevrei (Utterback).
Frierson, 1927, Check list N American naiades,
p. 95. Haas, 1969, Das Tierreich, pt. 88, p.
489.

Description. Shell rather small, seldom
reaching more than 40 mm in length. Out-
line elliptical, ovate or obovate. Valves in-
equilateral, solid, slightly inflated. An-
terior end regularly rounded; posterior end
of male rather pointed, of female more
broadly rounded. Ventral margin curved
in males, almost straight in females before
the marsupial swelling. Dorsal margin
slightly curved forming an indistinct angle
with the obliquely descending posterior
margin. Hinge ligament short. Posterior
ridge of the male, double, somewhat raised,
ending posteriorly in a sHght biangulation.
The ridges of the female fade out on the
marsupial swelling. Dorsal slope slightly
concave. The male has a rather wide, shal-
low, radial furrow, which ends in an emar-
gination. The medial ridge is only slightly
developed, and both it and the radial fur-
row are obscured in females by marsupial
swelling. Umbos moderately full and ele-
vated, located in the anterior third of the
shell; umbonal sculpture not observed. Sur-
face of the shell with irregular growth lines,
which are especially strong on the female
posteriorly. Periostracum rather shiny, yel-
lowish green, with numerous fine green
rays over the entire surface.

Left valve with two small, rough, sub-
compressed pseudocardinal teeth. No in-
terdentum. Two short, straight, lateral
teeth. Right xalve with one small, tri-
angular, pseudocardinal tooth, with a sec-
ond tiny tooth before it that is parallel to
the hinge line. One lateral tooth. Umbonal

Plagiola from Middle North America • Johnson

275

cavities shallow. Anterior adductor muscle
scars well impressed; posterior scars faint.
Pallial line distinct anteriorly. Nacre bluish
white. Shell heavier anteriorly, posterior
end of females especially thin and irrides-
cent.

Male shells are elliptical or ovate, with
a distinct raised double posterior ridge,
which ends in a biangulation near the base.
Before the ridge is a wide shallow radial
furrow and faint medial ridge.

The female is somewhat obovate, and
while the marsupial swelling obliterates the
radial furrow and the medial and posterior
ridges, there is a tendency for the shell to
be somewhat concave in the region of the
posterior ridges. The surface of the mar-
supial swelling is not different from the rest
of the shell, the feeble green rays are rather
evenlv distributed in both sexes.

Length Height Width

2.5

22
18

Holston River, Knox Co.,
Tennessee. Male.
Tennessee. Female. Hol-
otype of Lhiio deviatus
Anthony.

Remarks. Plagiola turgidula (Lea) most
closely resembles P. biemargitmta, but the
former has a more delicate, more elongate
shell. The male of turgidula differs from
that of hiemarginata, in that the former has
a less pronounced posterior and medial
ridge and the radial furrow is not as deep
as in the latter. Further, the posterior ridge
of turgidula ends higher on the posterior
margin than does that of biemarginata.

The female turgidula resembles the fe-
males of the other members of subgenus
Capsaefonnis, with its inflated, outcurved,
marsupial expansion in the area occupied
by the radial furrow in the male. However
in turgidula alone is there no color differ-
ence between the disk and the marsupial
swelling. Light green rays uniformly cover
the entire surface, except sometimes the ex-
treme anterior.

The male of this species was described
by Lea and the female, later, by Reeve.
Walker (1910: 81) was first to recognize

the two sexes as the same species. This is
the .species described by Utterback as lefev-
rei from the Ozark Plateau. He refers to
the male shell as having a post-umbonal
ridge, rather biangulated and with a very
slight radial furrow in front. P. turgidula
had been collected before 1914 on the
Ozark Plateau from Spring Creek, Hardy,
Sharp County, Arkansas, and so noted by
Simpson (1914; 1: 32).

The presence of P. turgidula, like that of
florentina on the Ozark Plateau, is evidence
of a former connection of that area with the
Cumberland Plateau.

Anatomij. Described by Utterback
(1916: 457 [192]).

Range, tapper White River system, Mis-
souri and Arkansas; Tennessee River sys-
tem, Tennessee and Alabama; Cumberland
River system.

Abundance. This species is known from
the Cumberland River, only on the authori-
ty of Lea and from a specimen, so labeled,
in the British Museum (Nat. Hist.). It
was once widely distributed throughout
the Tennessee River system, but according
to Stansbery (pers. comm. ), it is now re-
stricted to the Duck River, in the vicinity
of Normandy, Bedford County, Tennessee.

Listed a "extirpated" by Stansbery
(1976: 43, 51).

Specimens Examined
W^mTE River System

White River Drainage. Missouri: White
River, Moores Ferry [not located, but prob-
ably in Stone or Taney Co.] (MZUM).

Black River Drainage. Arkansas:
[Spring River], Hardy, Sharp Co. (MZUM;
MCZ, ex MZUM; Field Museum, Chicago).
Missouri: Black River, Williamsville,
Wayne Co. (Utterback).

Tennessee River System

Clinch River Drainage. Tennessee:
CHnch River (MCZ); Emory River, Harri-
man, Roane Co. (CM).

276 BuUctin Museum of Comparative Zoology, Vol. 148, No. 6

Holston River Drainage. Tennessee:
Holston River, Rogersville (MZUM); Hol-
ston River, Austin Mill (CM); both Haw-
kins Co.; mouth of Holston River, Austins
Crist Mill, Knox Co. (MCZ).

Elk River Drainage. Tennessee: Elk
River ( Stansbery, OSM).

Tennessee River Drainage. Alabama:
Shoals Creek (Hinkley and Marsh); Ten-
nessee River, Florence (Lea, USNM); botli
Lauderdale Co.

Bear Creek Drainage. Alabama: Bear
Creek, Burleson, Franklin Co. (CM).

Dnek River Drainage. Tennessee: Duck
River, Normandv (Stansbeiy, OSM); Shel-
byville (CM);'/;of/( Bedford Co. Duck
River, Columbia, Maury Co. (Hinkley and
Marsh).

Cumberland River System

Cumberland River Drainage. Cumber-
land River (Lea, USNM; BMNH).

Subgenus Pilea Simpson

Pilca Simpson 1900, Proc. U. S. Natl. Miis. 22: 522.
Type species, Unio pcisonatus, original designa-
tion.

Scalcnilla Ortmann and Walker 1922. Occ. Papers,
Mus. Zool., Univ. Mich. no. 112, p. 68. Type
species, Unio siilcatus Lea, original designa-
tion.

Obliquata Frierson, 1927, Check list N American
naiades, pp. 10, 52. Type species, Ohliquaria
(Scalenaria) ohliqitata Rafinesqne, original
designation, teste Errata et Corrigenda.

Description. Shell subtriangular or sub-
cjuadrate. "Male shell with a wide, shallow,
radiating depression in front of the pos-
terior ridge, that of the female with a
rounded, foliaceous swelling at the pos-
terior base. Animal with post basal flap of
mantle of female very heavy; ovisacs not
extending to the top of the marsupium."
( Simpson ) .

Remarks. Frierson (1927: 96) restricted
Simpson's subgenus Pilea to include per-
.sonatus and sulcatus {= obliquata), but
haysianus is now also included on the basis
of shell morphology. All three of these
.species tend to be proportionally higher in

relationship to length than any others in the
genus. They all have relatively heavy
shells, and have periostraca that are simi-
larly shiny or subshiny, greenish yellow,
or chestnut, with very fine green rays.
Unhke other members of Flagiola they
sometimes have purple or flesh-colored
nacre. The depression in front of the pos-
terior ridge in the male is a raised mar-
supial swelling in the female.

Key to the Species of Pilea

1. Shell snbtriangular, subquadrate, or sub-
orbicular 2

Shell subtrapezoid or subquadrate, nacre
white personata

2. Periostracum subshiny, yellowish brown or

greenish, nacre usually purplish obliquata

Periostracum shiny, tawny to chestnut, nacre
purplish haijsiana

Plagiola {Pilea) personata (Say)
Plate 14, figures 7-9
Distribution: Plate 6, figure B

Unio personatus Say 1829, New Harmony [Indi-
ana] Disseminator 2 ( 20 ) : 309 ( Wabash River,
[Indiana]; neotype MCZ 5763 from the Cum-
berland River, Tennessee, selected by Johnson
and Baker, 1973, Proc. Acad. Nat. Sci. Phila.
125: 166, pi. 8, fig. 5).

Unio pileiis Lea 1831, Trans. Amer. Philos. Soc.
4: 119, pi. 18, fig. 47 (Ohio River, near Cin-
cinnati [Hamilton Co.], Ohio, figured holotype
USNM 846()2a); 1834, Obs. Unio 1: 129.

Unio capilhiris Lea 1834, Trans. Amer. Philos. Soc.
5: 29, pi. 2, fig. 2 (Ohio, figured type [lost],
specimen subsequently identified by Lea USNM
84602); 18:34, Obs. Unio 1: 141.

Tiincilla (Pilca) personata (Say). Simpson, 1900,
Proc. U. S. Natl. Mus. 22: 522; 1914, Cat. Na-
iades 1: 23; 1903, Daniels, 27th Ann. Rept.
Dept. Geol. and Nat. Res. of Indiana, p. 646.

Dysnomia (Pilea) personata (Say). Ortmann,
1925, Amer. Mid. Nat. 9: 361. Frierson, 1927,
Check list N American naiades, p. 96, Haas,
1969, Das Tierreich, pt. 88, p. 480.

Dijsnomia personata (Say). Morrison, 1942, Bur.
Amer. Ethnology, Bull', no. 129, p. 364. Good-
rich and van der Schalie, 1944, Amer. Mid. Nat.
:32: 314. La Rocque, 1967, Geol. Surv. Ohio,
Bull. no. 62 (2): 278, fig. 169. Stansbery, 1970,
Malacologia 10: 19; 1971, Symposium of rare
and endangered moll. U. S., p. 18a, figs. 11, 12.

Description. Shell of medium size, reach-

Plagiola from Middle North America • Johnson 277

ing up to 65 mm in length. Outline subtri-
angular or subquadrate. Valves inequilat-
eral, solid, inflated. Anterior end regularly
rounded, sometimes subtruncate in the
male; posterior end of male regularly
rounded or subtruncate; posterior in fe-
males slightly produced above the median.
\^entral margin curv^ed with a slight emargi-
nation posteriorly in the male. Dorsal mar-
gin straight, forming an angle with the
obliquely descending posterior margin.
Hinge ligament short. Posterior ridge
faintly double in males, ending in a slight
bimargination, separated from a faint medi-
al ridge bv a narrow radial sulcus, which
ends in a slight emargination. The area of
the sulcus is a raised swelling marked by
denticulations in the female. Dorsal slope
slightly concave. Umbos full, somewhat
elevated and prominent, slightly turned
over a distinct lunule, located near the mid-
dle of the male shell and in the anterior
third of the female. Surface of the shell ir-
regularly concentrically sculptured. Perio-
stracum often silky or clothlike, greenish
yellow, brownish, or greenish brown in old
shells, usually with faint wavy green rays
over most of the surface.

Left \'alve with two chunky triangular
pseudocardinal teeth; slight interdentum.
Two short, almost straight, lateral teeth.
Right vah'c with one large triangular
pseudocardinal tooth, usually with a small
tooth before it, and sometimes with a much
smaller tooth after it. One lateral tooth,
sometimes with a vestigial tooth below.
Umbonal cavities shallow. Anterior and
posterior muscle scars well impressed.
Pallial line distinct anteriorly. Nacre white
or flesh colored. Shell heavier anteriorly,
marsupial area of the female quite thin.

Male shells are subtriangular with a dou-
ble posterior ridge that ends in a biangula-
tion in front of which is a narrow, shallow
radial depression that ends in a slight emar-
gination.

Female shells are subquadrate, the mar-
supial swelling occupies the place of the
radial depression. It is rather small, and

only slightly inflated, but it is radially
sculptured, denticulated, thin and exca-
vated within. Reaches the same size as the
male shell.

Length

Height

Width

111 111

Ohio River, Cincinnati,
Hamilton Co., Ohio.
Male.

As above. Female.

Habitat. Lives in deeper water [deeper
than what?] teste Morrison (1942: 364).

Remarks. Plagiola personata (Say)
most closely resembles P. ohliquata. Both
the male and the female of the former
have proportionally higher shells. The
radial furrow in the male personata is
more developed than is that of ohliquata.
The female of personata has a slightly
raised marsupial swelling, whereas that of
ohliquata is separated from the rest of the
shell by two distinct sulci.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem, Tennessee; Ohio River system, from
the lower Wabash drainage to Cincinnati,
Hamilton County, Ohio.

Ahundance. I know of no collections of
this species made in this century. It is pre-
sumed extinct." (Stansbery, 1976: 43, 50)

Specimens Examined
Tennessee River System

Clinch River Drainage. Tennessee:
CHnch River (MCZ, MZUM).

Tennessee River Drainage. Alahaina:
Tennessee River, Muscle Shoals, Colbert
and Lauderdale Cos. (CM); Tennessee
River, Florence, Lauderdale Co. (MCZ,
MZUM); Pickwick Basin, mounds be-
tween Barton, Colbert Co. and Waterloo,
Lauderdale Co. (Morrison, USNM).

Cumberland River System

Cumberland River Drainage. Tennes-
see: Cumberland River (MCZ); Cumber-
land River, near Priestly Shoals, Davidson
Co. (MZUM).

278 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Ohio River System

Wabash River Drainage. Indiana: White
River (MZUM); Wabash River (MCZ);
Wabash River, New Harmony, Posey Co.
( Daniels ) .

Ohio River Drainage. Ohio: Ohio
River, Cincinnati, Hamilton Co. (MCZ,
OSM, MZUM).

Plagiola {Piles) obliquata (Rafinesque)
Plate 14, figures 10-12
Distribution: Plate 4

Ohliquario obliquata Rafinesque 1820, Ann. Gen
des Sci. Physiques, Bruxelles 5: 309 (le Kentuky
[River]; lectotype ANSP 20226, selected by
Johnson and Baker, 1973, Proc. Acad. Nat. Sci.
Phila. 125: 163, pi. 7, fig. 1).

Unio sulcatus Lea 1829, Trans. Amer. Pliilos. Soc.
3: 430, pi. 9, fig. 12 (Ohio, figured holotype
USNM 84803); 1834, Obs. Unio 1: 44.

Unio ridibundus Say, 1829, New Harmony [Indi-
ana] Disseminator 2 (.5): 308; 1830, Amer.
Conch, no. 1, i^l. 5 (Cumberland River [Ten-
nessee], figured type [lost]).

Unio perplexus Say 1829, New Hannony [Indi-
ana] Disseminator 2 (.5): 309; 1830, American
Conch, no. 1 [letterpress to pi. 5] ( Cumberland
River [Tennessee], type [lost]), non Lea 1831.

Unio jlagellatus Say 1830, Amer. Conch, no. 1
[letterpress to pi. .5]. Listed as a synonym of
Unio sulcatus Lea.

Unio pcctitis Conrad 1853, Proc. Acad. Nat. Sci.
Phila. 6: 255 [nomen nudum]; 1854, Jour. Acad.
Nat. Sci. Phila. (2)2: 297, pi. 27, fig. 4 (Wa-
bash River [Indiana], type not located).

Unio stewardsonii stewensoni Gregorio 1914, II
Nat. Siciliano 22: 45, pl. 6, fig. 3 (no locality,
type presumed to be in Palenno Mus., Sicily
[not seen] ).

Unio propesulcatus Gregorio 1914, II Nat. Siciliano
22: 60, pl. 10, fig. 2 (Cumberland River, Ten-
nessee, type presumed to be in Palermo Mus.,
Sicily [not seen] ).

Truncilla (Scalenaria) sulcata (Lea). Simpson,
1900, Proc. U. S. Natl. Mus. 22: 520; 1914, Cat.
Naiades 1: 14.

Truncilla sulcata (Lea). Wilson and Clark, 1912,
U. S. Bur. Fisheries, Doc. 757: 31, 37, 38, 55.
Wilson and Clark, 1914, U. S. Bur. Fisheries,
Doc. 781: 46. Walker, 1918, Mus. Zool., Univ.
Mich., Misc. Pub. no. 8: 186.

Dysrwmia ( Scalenilla ) sulcata ( Lea ) . Ortmann
and Walker, 1922, Occ. Papers, Mus. Zool.,
Univ. Mich., no. 112, p. 68. Ortmann, 1925,
Amer. Mid. Nat. 9: 360. La Rocque, 1967,

Geol. Sur\-. Ohio, Bull. no. 62 (2): 278, fig.
171.

Dysnomia (Pilea) sulcata (Lea). Frierson 1927,
Check list N American naiades, p. 96. Haas,
Das Tierreich, pt. 88, p. 480.

Dysnomia sulcata (Lea). Morrison, 1942, Bur.
Amer. Ethnology, Bull. no. 129, p. 364. Good-
rich and van der SchaHe, 1944, Amer. Mid. Nat.
32: 314. Neel and Allen, 1964, Malacologia 1:
450. Stansbery, 1970, Malacologia 10: 19; 1971,
Symposium of rare and endangered moll. U. S.,
p. 18e, figs. 43, 44. Clark, 1977, Sterkiana,
nos. 65, 66, p. 27, fig. 19.

Dysnomia sulcata perobliquus Stansbery, 1970,
Malacologia 10: 19, non Conrad 1836.

Description. Shell of medium size, reach-
ing up to 70 mm in length. Outline sub-
trapezoid or quadrate. Valves very inequi-
lateral, inflated, solid. Anterior end of male
decidedly subtruncate, regularly rounded in
the female; posterior end of male somewhat
produced and pointed, truncate in the fe-
male. Ventral margin broadly curved in
males, almost straight in females. Dorsal
margin long and almost straight, forming
an angle with the obliquely descending
posterior margin. Hinge ligament rather
long. Posterior ridge of the male double,
rather low and faint, separated by a wide
faint radial furrow before another faint
ridge. Posterior ridge of female, somewhat
obscured before a shaip sulcus before the
marsupial swelling. Dorsal slope of male
very narrow, wider in the female. Umbos
full, elevated and prominent, turned over
a small lunule, located at the extreme an-
terior end, their sculpture consisting of a
few corrugations. Surface of the shell with
numerous strong growth rests. Periostracum
smooth and shiny, or subshiny, yellowish,
greenish yellow or brownish, usually feebly
rayed over the entire surface.

Left valve with two chunky triangular,
pseudocardinal teeth; slight interdentum.
Two short straight lateral teeth. Right valve
with one large triangular pseudocardinal
tooth, usually with a much smaller tooth
before and behind it. One lateral tooth,
sometimes with a vestigial tooth below. An-
terior and posterior muscle scars and pallial
line well impressed. Shell thinner posterior-

Plagiola from Middle North America • Johnson 279

ly in the female in the area of the mar-
supial swelling. Nacre purplish, pink, blu-
ish white or white.

Male shells are usually much larger than
those of the female, and are subtrapezoid.
The dorsal and ventral margins are curved,
but the anterior margin is subtnmcate. The
posterior end is pointed bluntly above, and
is slightly truncate below. The lower pos-
terior ridge is rather widely separated from
the medial ridge by a faint furrow.

Female shells are subquadrate, and tnm-
cated posteriorly. The marsupial area,
which occupies the space between the low-
er posterior and medial ridges in the male,
is inflated, rounded and separated from the
rest of the shell by two sulcations, the more
posterior being the more acute and ending
in a more acute emargination. The mar-
supial area is toothed and is sculptured by
foniier teeth at rest stops.

Length Height Width
mm mm mm

55 41 33

35 28 24

Ohio River, Cincinnati,
Hamilton Co., Ohio.
Male.

Cumberland River, Ten-
nessee. Female.

Remarks. Plagiola obliquata (Rafin-
esque) most closely resembles P. personata,
under which see: Remarks. The male shell
of personaia is subtriangular, while that of
obliquata is subtrapezoid. The umbos of
the latter project much farther forward
than do those of the male personata or of
the female obliquata, often reaching the
extreme anterior end as in Pleurobema clava
(Lamarck). The female of obliquata has
a marsupial swelling that is separated from
the rest of the shell by two distinct sulci.

Stansbery (1970: 19) regarded Unio gib-
bosus perobliquus Conrad as a geographi-
cally isolated subspecies from the streams
tributary to western Lake Erie and Lake
St. Clair on the basis of its having a wliite
nacre. Nacre color appears to be an eco-
phenotypic variation which changes from
purple to white throughout the range of P.
obliquata. The figured type of perobliquus
is P. torulosa ( Rafinesque ) .

Ortmann and Walker (1922: 68) pointed
out that the original description of obli-
quata applied as well to Pleurobema py-
ramidatum (Lea) as to Unio sulcatus Lea,
and made the reasonable suggestion that
obliquata be dropped as unidentifiable.
Frierson (1927: 53) argued that obliquata
was vyramidatum (Lea). He was followed
by Haas (1969: 297) and Morrison (1969:
24). Since the lectotype of obliquata is sul-
cata (Lea), the former taxon must, un-
fortimately, replace Lea's well-known
name.

Morrison (1969: 24) mentioned that
Unio sulcatus Lea 1829 is preoccupied, and
indicated that this taxon should be replaced
by Plagiola ridibundus (Say) 1829. Mor-
rison did not say by whom stdcatus was
preoccupied and a careful search of the
literature did not afford the answer, but
this is now of no interest here.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem, Kentuckv and Tennessee; Ohio River
system: Wabash River Drainage, and Ohio
River, to the Scioto River drainage, Ohio;
St. Lawrence River system: Lake Erie
drainage.

Abundance. As early as 1912, Wilson
and Clark (p. 31) were able to find only
one live specimen of P. obliquata in an ex-
tensive survey of the Maumee River drain-
age. Still occasionally found in streams trib-
utary to western Lake Erie or Lake St.
Clair (Stansbery, 1970: 19 as D. sulcata
perobliquus), and the Green River, Ken-
tucky (Stansbery, 1971: 18e). Epiobkisma
sulcata sulcata is listed as "extirpated" by
Stansbery (1976: 43, 50).

Specimens Examined

Tennessee Ri\^r System

Tennessee River Drainage. Alabama:
Tennessee River, Muscle Shoals, Lauder-
dale and Colbert Cos. (CM); Tennessee
River, Florence, Lauderdale Co. (MZUM);
Tennessee River, Tuscumbia. Colbert Co.
( MCZ); "Mound about 3 mi. WS^^' Gravel-
ly Springs, Lauderdale Co., only this speci-

280 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

men found in all mounds studied" (Morri-
son, 1942).

Cumberland River System

Cumherland River Drainage. Ken-
fiickij: Cumberland River, Rumside, Pu-
laski Co.; Cumberland Ri\'er, Neeleys Ford,
4 mi. S Rurks\ille, Cumberland Co.; {both
MZUM). Tennessee: Cumberland River,
Fort Blount, 6 mi. SW Jamesboro, Jackson
Co. (MZUM): Cumberland River, Coodall
Island, Smith Co. (W^ilson and Clark).

Canev Fork Drainage. Tennessee: Ca-
ney Fork, Buffalo Valley, Putnam Co. (Wil-
son and Clark ) .

Cumberland River Drainage. Tennes-
see: Cumberland River, Nashville, David-
son Co. (MCZ); Cumberland River, Half-
pone Bar, Cheatham Co. (Wilson and
Clark).

Harpeth River Drainage. Tennessee:
Harpeth River (MZUM).

Ohio River System

Ohio River Drainage. Illinois: Ohio
River ( Baker).

Wahash River Drainage. Indiana:
\Miite River (MCZ); West Fork, White
Rixer ( Daniels ) ; White River, Indianapolis
(MZUM); both Marion Co.; White River,
Rockford, Jackson Co. (Daniels); Wabash
River, Lafayette, Tippecanoe Co. (MCZ
and MZUM); Wabash River, New Har-
mony, Posey Co. (MZUM).

Green River Drainage. Kentucky:
Green River, Glenmore, ^^■arren Co.
(OSM); Green River, Rochester, Butler Co.
(MCZ; MZUM).

Kentucky River Drainage. Kentucky:
Kentucky River (Rafinesque).

Ohio River Drainage. Ohio: Ohio
River, Cincinnati, Plamilton Co. (MCZ).

Licking River Drainage. Kentucky:
Licking River (MCZ).

Scioto River Drainage. Ohio: Scioto
River (MZUM).

St. Lavh^ence River System

Great Lakes Drainage. (Lake Erie)

Ohio: Blanchard River, Findlay, Hancock
Co. (MZUM); Fish Creek, Williams Co.
(Clark, 1977: 21). hidiam: St. Joseph
River (MZUM); St. Marys River (Wil-
son and Clark); Maumee River (MCZ,
MZUM); both Fort Wayne, all Allen Co.
Ohio: Maumee River, 4 mi. below Defi-
ance; Auglaise River, 4 mi. above Defiance;
both Defiance Co. (both Wilson and
Clark). Lake Erie, Putin Bay, Ottawa Co.
(Wilson and Clark). Michigan: Otter
Creek, Monroe Co. (MZUM). Lake St.
Clair (Stansbery, 1970). Detroit River,
Belle Island, Wayne Co. (MZUM). On-
tario: Detroit River, Bois Blanc Isle, Essex
Co. (MZUM). Michigan: Lake Erie, near
Stony Creek; River Rasin; Lake Erie, La
Plaisance Bay; all Monroe Co. (all MZUM).
New York: Niagara River (MZUM).

Ptagiola (Pilea) haysiana (Lea)
Plate 15, figures 1-4
Distribution: Plate 8, figure A

Vnio haysianus Lea 1834, Trans. Anier. Philos.
Soc. 5: 35, pi. 3, fig. 7 (Cumberland River
[Tennessee], figured type [lost], male speci-
men, subsequently identified by Lea, USNM
84613 from the Cumberland River, Nashville,
Davidson Co., Tennessee); 1834, Obs. Unio 1:
147.

Unio sowerhyanus Lea 1834, Trans. Amer. Philos.
Soc. 5: 68, pi. 10, fig. 28 (Tennessee, figured
type [lost], 4 male specimens, subsequently
identified by Lea, MCZ 178886 from the Caney
Fork of the Cumberland Ri\er, Tennessee);
1834, Obs. Unio 1: 180.

Truncilla (Scalcnaria) haysiana (Lea). 1900,
Proc. U. S. Natl. Mus. 22: 520; 1914, Cat. Na-
iades 1: 16.

Truncilla haysiana (Lea). Wilson and Clark,
1914, U. S. Bur. Fish. Doc. no. 781, p. 46. Ort-
mann, 1912, Ann. Carnegie Mus. 8: 357; 1913,
Proc. Amer. Philos. Soc. 52: 311; 1918, Proc.
Amer. Philos. Soc. 57: 587.

Dysnomia (Scalenilla) haysiana (Lea). Ortmann,
1925, Amer. Mid. Nat.' 9: 361.

Dysnomia (Pcnita) haysiana (Lea). Frierson,
1927, Check list N American naiades, p. 94.
Haas, 1969, Das Tierreich, pt. 88, p. 483.

Dysnomia haysiana (Lea). Morrison, 1942, Bur.
Amer. Ethnologv, Bull. no. 129, p. 364. Neel
and Allen, 1964, Malacologia 1: 450, fig. 60.
Stansbery 1970, Malacologia 10: 19; 1971, Sym-
posiimi of rare and endangered moll. U. S., p.
18e, figs. 45, 46.

Plagiola from Middle North America • Johnson 281

Epiohlasma haysiana (Lea). Stansberv, 1972,
Anier. Mai. Union, Bull, for 1972, p. 22.

Description. Shell usually of small size,
reaching up to 40 mm in length. Outline
subtriangular or suborbicular. Valves in-
equilateral, somewhat inflated, solid. An-
terior end regularly rounded, posterior end
of male somewhat elongate and slightly
more broadly rounded; more broadly
rounded in the female. Ventral margin
curved. Dorsal margin curved merging al-
most imperceptibly with the obliquely de-
scending posterior margin. Hinge ligament
short. Posterior ridge of the male faint, but
double, broadly curved or flat, merging
into a flat dorsal slope; ridge elevated into
a marsupial swelling in females. There is
a broad, shallow radial furrow before a
medial ridge in the male; the furrow is
narrow and deep in the female. The pos-
terior end of the male shows vestiges of
teeth along the growth rests; these are
prominent in the female especially on the
marsupial sweUing. Umbos full and high,
located somewhat anteriorly, feebly sculp-
tured. Surface of the shell smooth anteri-
orlv, but sometimes rendered subnodulous
by deep growth rests. Periostracum very
shiny, especially anteriorly, tawny to chest-
nut, with a few greenish rays in the radial
fiurow and on the disk.

Left \al\'e with two chunky, triangular
pseudocardinal teeth of about equal size
with a sharp, deep, triangular pit between
them extending to the hinge line; inter-
dentum short but wide. Two short straight
lateral teeth. Right valve with one large,
triangular, pseudocardinal tooth, sometimes
with a \'estigial tooth anterior to it. One
well-developed lateral tooth, with a vesti-
gial tooth below. Umbonal cavities shallow.
Anterior and posterior muscle scars and
pallial line well impressed. Marsupial area
of the female thinner and somewhat ex-
cavated. Nacre usually purple, but some-
times white and iridescent posteriorly.

Male shells are subtriangular, often as
high or higher than long. The radial fur-
row in front of the posterior ridge is broad

and shallow. The posterior end has ves-
tiges of teeth along the growth lines.

Female shells are suborbicular, less long
than high. The posterior ridge is inflated,
and extends below the ventral margin. The
radial furrow in front of the posterior ridge
is narrow, sharp and deep. The posterior
end has more vestiges of sharper teeth
along the growth lines than does the male.

Length Height Width
mm mm mm

40 31 21

Clinch River, Union Co.,
Tennessee. Male.
As above. Female.

Anatomy. Discussed by Ortmann (1912,
357).

Remarks. Plagiola haysiana (Lea) is
easily distinguished from the other mem-
bers of the genus by its polished, tawny to
chestnut periostracum, its small size, its un-
usually thick and heavy shell, and its usu-
ally purplish nacre.

Lea described the female of tliis species
as U. haysianus and the male as U. sower-
byanus.

Range. Tennessee River system, Vir-
ginia, Tennessee and Alabama; Cumberland
River system, Kentucky and Tennessee.

Abundance. Now restricted to the Clinch
River between St. Paul, Wise County, to
Dungannon, Scott County, \^irginia, a dis-
tance of only about 10 miles (Stansbery,
1970: 19). Listed as "extirpated" bv Stans-
bery ( 1976: 43, 50).

Specimens Examined

Tennessee River System

Powell River Drainage. Virginia: Pow-
ell River, 2.5 mi. S Jonesville; Wallens
Creek; Powell River, Lyttons Mill (all
MCZ); Powell Ri\'er, Pennington Gap
( CM ) ; all Lee Co.

Clinch River Drainage. Virginia: Clinch
River, Raven, Tazewell Co. (CM); Clinch
River, Cleveland, Russell Co. (MCZ);
Clinch River, Saint Paul, Wise Co. (Stans-
bery, OSM); Clinch River, Dungannon,
Scott Co. (MCZ). Tennessee: Clinch

282 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

River, Union Co.; Clinch River, Clinton,
Anderson Co. (both MCZ).

Holston River Drainage. Virginia: North
Fork, Hilton, Scott Co. (CM). Tennessee:
Sonth Fork, Pactolus, Sullivan Co. (CM);
Mouth of Holston Ri\'er, Austins Grist Mill,
Knox Co. (MCZ).

Tennessee River Drainage. Tennessee:
Tennessee River, near Knoxville, Knox Co.
(MCZ).

Little River Drainage. Tennessee: mouth
of Little River, Little River Shoals, Blount
Co. (MCZ).

Little Tennessee River Drainage. Ten-
nessee: Little Tennessee River, Monroe Co.
(MCZ); Little Tennessee River, Coytee,
Loudon Co. (MZUM).

Elk River Drainage. Tennessee: Elk
River, Fayetteville, Lincoln Co. (MCZ).
Alabama: Lower Elk River (Conrad).

Tennessee River Drainage. Alabama:
Tennessee River, Florence, Lauderdale Co.;
Tennessee River, Tuscunibia, Colbert Co.
{both MCZ); Pickwick Basin, mounds be-
tween Barton, Colbert Co. and Waterloo,
Lauderdale Co. (Momson, USNM).

Cumberland River System

Cumberland River Drainage. Ken-
tucky : Big South Fork, above Burnside and
Parkers Lake Station; both Pulaski Co.
{J)Oth MCZ). Tennessee: Cumberland
River, Goodall Island, Smith Co. (Wilson
and Clark ) .

Canev Fork Drainage. Tennessee: Ca-
ney Fork, [Smith Co.] (MCZ).

Cumberland River Drainage. Tennes-
see: Cumberland River, Nashville, David-
son Co. (MCZ; USNM); Cumberland Riv-
er, Clarksville, Montgomery Co. (Wilson
and Clark ) .

Subgenus Epioblasma Rafinesque

Epiohlasma Rafinesque 1831, Cont. Monog. Bi-
valve Shells River Ohio, p. 2. Type species,
Epiohlasma biloba Rafinescjue, monotypic.

Dymornia Agassiz 1852, Arch. Hir Naturgesch. 18:
43. First species listed, [Ohliquaria] jlexuosa
[Rafinesque] = Vnio foliaftts Hildreth [teste

Agassiz]. Type species, Unio foliatus Hildreth,
subsequent designation, Simpson, 1900, Proc.
U. S. Natl. Mus. 22: 521.

Description. Shell subrhomboid, sub-
quadrate or subtriangular. "Shell of the
male with a posterior and central radiating
ridge, with a wide, flattened space be-
tween them; that of the female with a
greatly produced inflation, which is but a
little behind the center of the base and
which is a continuation of the central
ridge" (Simpson, 1914: 18).

Renuirks. The availabihty of Epioblasma
over Dysnomia becomes of nugatory in-
terest once both of these names are reduced
to subgeneric standing under Plagiola. The
availability of Epioblasma is dependent on
the identification of £. bioloba. Frierson
(1914: 7) asserted that E. bioloba Raf. =
U. foliatus Hild. [=flexuosa Raf.]. In reply
to Frierson, Ortmann and Walker (1922:
71) gave a number of palpable reasons why
bioloba was not recognizable to them. In
the index to his Check list of North Ameri-
can naiades, Frierson (1927: 101) indicated
that bioloba belonged in the synonymy of
Dysnomia flexuosa (Raf.) although he ne-
glected to include it in the synomymy of
the latter. He apparently forgot he had
previously asserted the availability of Epio-
blasma over Dysnomia. Thiele (1934: 837),
Clench ( 1959:' 1157), Monison (1969: 24),
and Stansbery (1972: 22) recognize Epio-
blasma. The latter said (1973, pers.
comm.), "£pio/;/«.s7?w2 [bioloba] Rafinesque
1831 is clearly the female of [Obliquaria]
flexuosa Rafinesque 1820."

Van der Schalie (1973: 49) partially re-
quoted Ortmann and Walker as to why
bioloba is unrecognizable, merely reassert-
ing their position. While no one has made
any contribution to the subject of bioloba' s
recognizability since Ortmann and Walker,
most subsequent published opinions, as
noted above, are that bioloba is identifiable
and is the female of flexuosa.

The type and an authentic specimen sent
to Fenissac of bioloba are both lost (John-
son and Baker, 1973: 149), therefore in the

Plagiola from Middle North America • Johnson 283

interests of stability of nomenclature, the
"exceptional circumstances" [Int. Code
Zool. Nomen., 1964, Art. 75 (a) (i)] de-
scribed above require that a neotype be
selected for hioJoha. This is done under
Flagioh {Epiohlasma) flexuosa (Rafin-
esque ) .

Key to the Species of Epioblasma

Medial ridge considerably elevated flexuosa

Medial ridge not much elevated stewardsoni

Plagiola {Epioblasma) flexuosa
(Rafinesque)

Plate 15, figures 5-8

Distribution: Plate 5

Ohliquaria flexuosa Rafinesque 1820, Ann. Gen.
Sci. Physiques, Bruxelles 5: 306 (le Kentuky,
Salt-ri\er et Green-river; lectotype ANSP 20249
from the Kentucky River, selected by Johnson
and Baker, 1973, 'Proc. Acad. Nat. Sci. Phila.
125: 163, pi. 7, fig. 2).

Vnio foliatus Hildreth 1828, American Jour. Sci.
14: 284, fig. 16 (Ohio; known only from the
figured type [lost] ).

Epioblasma biloba Rafinesque 1831, Cont. Monog.
Bivalve Shells River Ohio, p. 2 (Green River
and Kentucky River; type lost, teste Johnson and
Baker, 1973, Proc. Acad. Nat. Sci. Phila. 125:
149). Neotype, here selected, ANSP 56571, pi.
15, fig. 7, and the type locality restricted to the
Ohio River, near Cincinnati, Hamilton Co.,
Ohio).

TnmciUa lewisi Walker 1910, Nautilus 24: 42, pi.
3, fig. 3 female; figs. 4, 5 male ( Holston River,
Tennessee, the figured female svntype is, here
selected, lectotype MZUM 91456^ pi. 15, fig. 8).
Simpson, 1914, Cat. Naiades 1: 20. Ortniann,
1918, Proc. Amer. Philos. Soc. 57: 588.

Tmncilla foliata (Hildreth). Simpson, 1900, Proc.
U. S. Natl. Mus. 22: 521; 1914, Cat. Naiades 1:
18.

Dysnomia (Dtjsuomia) flexuosa (Rafinesque).
Ortmann and Walker, 1922, Occ. Papers, Mus.
Zool. Univ. Michigan no. 122, p. 70. Ortmann,

1926, Ann. Carnegie Mus. 17: 183. Frierson,

1927, Check list N American naiades, p. 93.
Goodrich and \an der Schalie, 1944, Amer. Mid.
Nat. 32: 314. La Rocque 1967, Geol. Surv.
Ohio, Bull. 62 (2): 275, fig. 167. Haas, 1969,
Das Tierreich, pt. 88, p. 478. Stansbery, 1970,
Malacologia 10: 19, pi. 1, figs. 1, 2; 1971, Sym-
posium of rare and endangered moll. U. S., p.
18a, figs. 5, 6.

Dysnomia (Dysnomia) lewisi (Walker). Frierson,

1927, Check list N American naiades, p. 93.

Haas, 1969, Das Tierreich, pt. 88, p. 479.
Dysnomia flexuosa lewisi (Walker). Morrison,

1942, Bur. Amer. Ethnology, Bull. no. 129, p.

366.
Dysnomia lewisi (Walker). Neel and Allen, 1964,

Malacologia 1: 450, figs. 61, 66. Stansbery 1970,

Malacologia 10: 19; 1971, Symposium of rare

and endangered moll. U. S., p. 18a, fig. 7 [fig.

8, after Walker, is a male of E. stewardsoni].

Description. Shell of medium size,
reaching up to 75 mm in length. Outline
of male subrhomboid or quadrate; outline
of female, depending on the extent of mar-
supial swelling, subtriangular. Valves equi-
lateral, slightly inflated, solid. Anterior end
of the male shell regularly rounded, forming
an obtuse angle at its junction with the
basal emargination. Anterior end of the fe-
male regularly rounded, becoming straight
and obliquely descending beyond the base
line. Posterior end of the male subtrun-
cated; that of the female somewhat ex-
tended and pointed. Ventral margin of the
male with two slight emarginations; one
considerable emargination in females. Dor-
sal margin broadly curved in males; margin
short and straight in females, forming an
obtuse angle with the obliquely descending
posterior margin. Hinge ligament promi-
nent. Posterior ridge distinct, rounded to-
ward the imibos, becoming flattened and
somewhat double as it approaches the pos-
terior end, where it terminates in a slight
biangulation projecting a little beyond the
posterior and basal lines. Dorsal slope nar-
row and slightly concave. In front of a
median groove is a strong anterior ridge
that becomes more pronounced as it ap-
proaches the base, where it terminates in an
angle sHghtly backward, at, or a little be-
hind, the middle of the base. In the female
this anterior ridge becomes an enormously
produced lobe, or winglike marsupial
swelling. Umbos laterally compressed, only
slightly elevated above the hinge line, lo-
cated near the middle of the shell, their
sculpture faint and cormgated. Surface of
the shell with uneven concentric sculpture.
Periostracum uniformK' pale brownish
green or brownish, ^\'ith faint green rays.

284 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

except for the female's marsupial swelling,
which may be dark green.

Left valve with two ragged pseudocardi-
nal teeth, triangular and of about equal
size; the anterior tooth narrow, straight, di-
rected oblicjuely forward, slightly widening
toward the anterior end; the posterior tooth
triangular; the space between them tri-
angular and extending to the hinge. Inter-
dentum rather long, narrow, rounded and
parallel to the hinge. Two nearly straight,
granular, lateral teeth, bent obliquely down-
ward from the hinge line. Right valve with
two pseudocardinal teeth, the anterior tooth
small but well developed; the posterior
tooth long and triangular, separated from
the interdentuni by a deep groove. One
well developed lateral tooth, often with a
parallel vestigial tooth below. Umbonal
cavities rather shallow. Anterior and pos-
terior muscle scars well impressed. Pallial
line distinct. Nacre white.

Although the wide radial furrow of the
male is usually a little deeper than in the
female, the shells of the two sexes are
essentially alike until they are about one-
third grown. Then the female begins to de-
velop a rounded, prolonged marsupial swell-
ing, which points backward, almost at the
middle of the ventral margin. The swelling
is scarcely, if at all, radially sculptured, but
it is thin, winglike and gapes slightly.

Wabash River, Indiana.
Male.

As above. Female.
Holston River, Austins
Grist Mill, K,nox Co.,
Tennessee. Male.
Holston River, Tennessee.
Female. Lectotype of
Truncilla lewisi.

Anatomy. Known only from a brief de-
scription, based on an imperfect dry speci-
men, by Lea ( 1863, Jour. Acad. Nat. Sci.
Phila. (2) 5: 443; 1863, Obs. Unio 10: 79).

Habitat. Call (1898: 511) observed that
in the Ohio River this species was found
on muddy bottoms in deeper water, while
Stansbery (1970: 19) suggested that jlexu-

.ength
mm

Height
mm

Width
mm
41

76
37

68
30

49.5

osa was, "apparently a species of shallow
riffles in big rivers," which it was in the
Tennessee and Cumberland River systems.

Remarks. The pronounced sexual di-
morphism shown in the shells of Phiffiola
flexuosa ( Rafines(iue ) , reflecting the spe-
cial use of the gills as marsupia, make it
one of the world's more remarkable species
of Unionacea. It cannot be confused with
any other member of the genus save P. (£.)
steicarchoni, under which see Remarks.
Walker described the form of flexuosa
found in the Tennessee and Cumberland
river systems as lewisi on the basis that the
male of the latter did not grow as large or
have a shell as heavy as that of the former.
The female, he said, 'l^esides being uni-
formly smaller, more delicate and smoother
than [flexuosa], is especially characterized
by the difference in the marsupial expan-
sion, which is triangular and comparatively
narro\\' at the extremity and of a different
texture from the body of the shell [being
thin and dark green]. In [flexuosa] this ex-
pansion is broadly rounded and is of the
same texture as the remainder of the valve."

Truncilla leuisi is regarded here as
merely an ecophenotypic variant. Plagiola
flexuosa does not grow as large in the Ten-
nessee and Cumberland river systems,
where it lives on shallow riffles. It seems
to have found in the muddy Ohio River,
where it spread in post-glacial time, a favor-
able environment where it could achieve
larger size. While Ohio River females do
not usually have green marsupial swellings,
one in the Museum of Comparative Zo-
ologv, no. 5358, does have the green mar-
supial swelling and is indistinguishable
from females from the Tennessee and Cum-
berland river systems.

Specimens of Pla<^iola flexuosa. Museum
of Comparative Zoology 221711, from the,
"Ohio River, Stubenville, Ohio, C. M.
Wheatley, 1856," and so labeled in the
shells, extend the range of this species far-
ther up the Ohio than was previously au-
thenticated. While no specimens of this
species have been found in the Kentucky,

Plagiola from Middle North America • Johnson 285

Salt and Green rivers, since it was reported
from them by Rafinesque, these localities
fall within the known range of the species.

Range. Tennessee River system, Tennes-
see and Alabama; Cumberland River sys-
tem, Kentucky; Ohio River system; from
the lower \\'abash River, Indiana to the
Ohio River, Jefferson Co., Ohio.

Abundance. No collection has very many
examples of this rare shell. Found sparingly
in the Pickwick Basin mound deposits by
Morrison (1942: 366). "Recorded from
both the Tennessee and Cumberland River
systems up until the construction of \\'olf
Creek Dam on the Cumberland [collected
by Neel and Allen 1947-49] and the TVA
Dams on the Tennessee. It has not been
collected in over 20 years and hence is pre-
sumed extinct." ( Stansbery, 1970, 19 as
lewisi). Once locally abundant in the Ohio
River, near Cincinnati, Hamilton County,
Ohio. "This species has not been collected
since 1900 [in the Ohio River] and is pre-
sumed extinct" (Stansbery, 1970: 19).
Both lewisi and fle.xuosa are listed as "ex-
tirpated" by Stansbery (1976: 43, 50).

Speceniexs Examined
Tennessee River System

Powell River Drainage. Tennessee:
Powell River, Combs, Claiborne Co. (CM,
single male ) .

Clinch River Drainage. Tennessee:
ClinchRiver (MCZ).

Holston River Drainage. Tennessee:
Holston River, Holston Station, Grainger
Co. (CM, single male); mouth of Holston
River, Austins Grist Mill, Knox Co. (MCZ).

Tennessee River Drainage. Alabama:
Tennessee River, Tuscumbia, Colbert Co.
(USNM); Pickwick Basin, mounds between
Barton, Colbert Co. and Waterloo, Lauder-
dale Co. (Morrison, USNM).

Cumberland Ri\ter System

Cumberland River Drainage. Ken-
tucky: Cumberland River, Port Burnside,
Pulaski Co. (MZUM); Cumberland River,

Rowena Ferry; Cumberland River, Long
Bottom, just below Wolf Creek Dam, both
Russell Co. {both MZUM).

Ohio River System

Wabash River Drainage. Indiana:
White River (USNM); Wabash River
(MCZ); Wabash River, New Harmony,
Posey Co. (MZUM).

Green River Drainage. Kentucky: Green
River ( Rafinesque ) .

Salt River Drainage. Kentucky: Salt
River ( Rafinesque ) .

Kentucky River Drainage. Kentucky:
Kentucky River ( Rafinesque ) .

Ohio River Drainage. Ohio: Ohio
River, Cincinnati, Hamilton Co. (MCZ).
Kentucky: Fort Ancient Aspect, Campbell
Co. [from Indian Midden] (USNM). Ohio:
Ohio River, Steubenville, Jefferson Co.
(MCZ).

Plagiola {Epioblasma) stewardsoni (Lea)
Plate 15, figures 9, 10
Distribution: Plate 8, figure C

Unio steivardsoni Lea, 1852, Trans. Amer. Philos.
Soc. 10: 278, pi. 23, fig. 36 ([Tennessee] River,
Chattanooga [Hamilton Co.], Tennessee; figured
holotvpe ANSP 56572); 1852, Obs. Unio 5: 34.

Truncilla stewardsoni (Lea). Simpson, 1900,
Proc. U. S. Natl. Mns. 22: 521; 1914, Cat. Na-
iades 1: 21. Ortmann, 1918, Proc. Amer. Philos.
Soc. 57: 588.

Dijsnomia (Dysuomia) stewardsoni (Lea). Ort-
mann, 1925, Amer. Mid. Nat. 32: 364. Frier-
son, 1927 Check list N American naiades, p. 93.
Haas, 1969, Das Tierreich, pt. 88, p. 478.

Dijsnomia stewardsoni (Lea). Morrison, 1942,
Bur. Amer. Ethnology, Bull. no. 129, p. 365.
Stansbery, 1970, Malacologia 10: 19, pi. 1, figs.
3, 4; 1971, Symposium of rare and endangered
moll. U. S., p. 18a, figs. [8 as male of lewisi]
9, 10; Hurd, 1974, Ph.D. thesis, p. 97.

Description. Shell of small size, usually
not exceeding 45 mm in length. Outline of
shell irregularly rhomboid. \'ahes subequi-
lateral, subcompressed, solid. Anterior end
of the male shell regularh- rounded forming
an obtuse angle at its junction with the
basal emargination. Anterior end of the fe-
male regularh^ rounded but becoming

286 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

straight and obliquely descending some-
what beyond the base line. Posterior end
of the male subtruncated; that of the fe-
male slightly extended. Ventral margin of
the male emarginate; female margin more
considerably emarginate. Dorsal margin
broadly cnrs^ed in males; short and straight
in females, forming an obtuse angle with
the obliquely descending posterior margin.
Hinge ligament prominent. Posterior ridge
distinct and rounded, becoming slightly bi-
angulate as it approaches the posterior end,
where it terminates in a slight projection.
Dorsal slope nanow and slightly concave.
In front of a median groove there is a
rounded anterior ridge that becomes some-
what pronounced as it approaches the base,
where it terminates in a slight angle slightly
backward, at, or a little behind, the mid-
dle of the ventral margin. In the female this
anterior ridge becomes a produced, broadly
rounded marsupial swelling. Umbos later-
ally compressed, only slightly elevated
above the hinge line, located near the mid-
dle of the shell, their sculpture not ob-
served. Surface of the shell with uneven
concentric sculpture. Periostracum green-
ish yellow to brownish, with faint green
rays. Marsupial expansion of the female
sometimes dark and greenish.

Left valve with two ragged pseudo-
cardinal teeth, triangular, and of about
equal size; the anterior tooth narrow,
straight, directed obliquely forward, slightly
widening toward the anterior end; the pos-
terior tooth triangular; the space between
them triangular and extending to the hinge.
Interdentum rather long, narrow, rounded,
and parallel to the hinge. Two nearly
straiglit, granular, lateral teeth, bent
obli(|uely down from the hinge line. Right
valve with two pseudocardinal teeth, the
anterior tooth small and very slightly ele-
vated above the hinge line, the posterior
tooth long, high and triangular, separated
from the interdentiun by a groove. One
well-developed lateral tooth, often with a
parallel vestigial tooth below. Umbonal
cavities rather shallow. Anterior and pos-

terior adductor muscle scars well impressed.
Pallial line distinct in males, distinct an-
teriorly only in females. Nacre white.

Although the wide radial furrow of the
male being a very little bit deeper than in
the female, the shells of the two sexes are
essentially alike until they are about two-
thirds grown. Then the female begins to
develop a rounded, prolonged marsupial
swelling which points backward, almost at
the middle of the base. The swelling is of-
ten faintly radially sculptured, and is thin
and winglike.

Length Height Width
mm mm mm

40 28 16

32
31

27
29

Holston River, Knoxville,
Knox Co., Tennessee.
Male.

As above. Female.

[Tennessee] River, Chat-
tanooga [Hamilton Co.],
Tennessee. Female. Holo-
type of U. stewardsoni
Lea.

Remarks. Plagiola stewardsoni (Lea)
and flexiiosa are clearly sibling species.
The former differs from flexuosa in several
ways: It does not attain as large a size, the
male shell is more quadrate, and the radial
furrow is not as distinct or as deep. In the
female of steioardsoni, the radial furrow is
quite feeble and runs into the marsupial
swelling, where it is generally obliterated,
while in flexuosa the furrow continues to
the base of the shell behind the marsupial
swelling. The posterior ridge of steicard-
.soni is not produced posteriorly, which ren-
ders the ventral emargination proportion-
ally more acute than in flexuosa. The
marsupial swelling of .stewardsoni is not as
exaggerated as in flexuosa, but it also tends
to be darker, often greenish, as in flexuosa
specimens from the Tennessee and Cumber-
land river systems.

The two records listed by Hurd (1974:
97, 116) from the Coosa River drainage of
the Mobile-Alabama-Coosa river system:
"Coosa River, Al." (MZUM 90564) and
"Etowah River, Ga." (MZUM 90565), as

Plagiola from Middle North America • Johnson 287

suggested to Hurd by van der Schalie, are
doubtless spurious.

Ranis^e. Restricted to the Tennessee and
Cumberland river systems, Tennessee and
Alabama.

Ahiinclonce. A rare species that has never
been found in great numbers. This fact was
noted b>' Ortmann (191S: 588) and by
Morrison (1942: 365) who found it spar-
ingly in the Pickwick Basin mound sam-
ples. "A rare species even before the im-
poundments and apparently not collected
in the last half century. It is presumed ex-
tinct" (Stansbeiy, 1976: 43, 50).

Specimens Examined
Tennessee River System

Clinch River Drainage. Tennessee:
Clinch Ri\'er (MCZ); Clinch River, Clinton
(CM); J)Oth Anderson Co.

Holston River Drainage. Tennessee:
Holston River, Holston Station, Grainger
Co.; Holston River, Mascot; Holston River,
McMillan {all CM); mouth of Holston
River, Austins Grist Mill (MCZ); all Knox
Co.

French Broad River Drainage. Tennes-
see: Nolichucky River (MCZ).

Tennessee River Drainage. Tennessee:
Tennessee River, Knoxville, Knox Co.
(MCZ).

Little River Drainage. Tennessee:
mouth of Little River, Little River Shoals,
Blount Co. (MCZ).

Tennessee River Drainage. Tennessee:
Tennessee River, Chattanooga, Hamilton
Co. (Lea, USNM; MZUM). Alabama:
Tennessee River, Bridgeport, Jackson Co.
(MCZ); Tennessee River, Florence, Lau-
derdale Co.; Tennessee River, Tuscumbia,
Colbert Co. (both USNM); Pickwick Ba-
sin, mounds between Barton, Colbert Co.
and ^^■ aterloo, Lauderdale Co. ( Morrison,
USNM).

Cumberland River System

Cunil>erland River Drainage. Ken-
tucky: Cumberland River, Pulaski Co.

(MZUM). Tennessee: Cumberland River
(USNM); Bartons Creek, Lebanon, Wilson
Co. (MCZ).

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are repaged, but the plate and figure num-
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Meyer, E. R. 1974. Unionid mussels of the Wa-
bash, White and East Fork of the White riv-
ers, Indiana. Virginia Jour. Sci. 25 ( 1 ) : 20-
25.

Morrison, J. P. E. 1942. Preliminary report on
mollusks found in shell mounds of the Pick-
wick Landing Basin in the Tennessee River
Valley. Bureau Amer. Ethnology (Smithson-
ian Inst., Washington) Bull. no. 129: 339-
392.

. 1969. The earliest names for North

American naiades. Amer. Mai. Union, Repts.
for 1969, pp. 22-24.

Murray, H. A. and B. Leonard. 1962. Unionid
mussels in Kansas. Univ. of Kansas, Mus.
Nat. Hist. Misc. Pub. no. 28, pp. 1-184, 45
pis., 42 figs.

Neel, J. K. and W. R. Allen. 1964. The mus-
sel fauna of the upper Cumberland Basin be-

Plagiola from Middle North America • Johnson 289

fore its impoundment. Malacologia 1 ( 3 ) :
427-459, figs.

Ortmann, a. E. 1909. A preliminary list of
the Unionidae of western Pennsylvania, with
new localities for species from eastern Penn-
sylvania. Ann. Carnegie Mus. 5 (2-3): 179-
210.

. 1911. A monograph of the na jades of

Pennsylvania. Part 1. Anatomical investiga-
tions. Part 2. The system of North Ameri-
can Najades. Mem. Carnegie Mus. 4 (6):
279-347, pis. 86-89.

. 1912a. Notes upon the families and

genera of najades. Ann. Carnegie Mus. 8
( 2 ) : 222-365, pis. 18-20.

. 1912b. The geological origin of the

freshwater fauna of Pennsylvania. Pennsyl-
vania Topog. and Geol. Survey 1910-1912,
pp. 130-149, fig. 1.

. 1913. The Alleghenian Divide, and its

influence upon the freshwater fauna. Proc.
Amer. Philos. Soc. 52: 287-390, pis. 12-14.

. 1918. The Nayades (freshwater mus-
sels) of the upper Tennessee drainage. With
notes on synonymy and distribution. Proc.
Amer. Philos. Soc. 57 (6): 521-625.

. 1919. A monograph of the naiades of

Pennsylvania. Part 3. Systematic account of
the genera and species. Mem. Carnegie Mus.
8(1): 1-384, pis. 1-21.

. 1920. Correlation of shape and station

in freshwater mussels ( Naiades ) . Proc. Amer.
Philos. Soc. 59 (4): 269-312.

. 1924a. The naiad-fauna of Duck River

in Tennessee. Amer. Mid. Nat. 9(1): 18-62.

— . 1924b. Distributional features of naiades
in tributaries of Lake Erie. Amer. Mid. Nat.
9: 101-117.

— . 1925. The naiad-fauna of the Tennes-
see River system below Walden Gorge. Amer.
Mid. Nat. 9 (8): 321-373.

. 1926. The naiades of the Green River

drainage in Kentucky. Ann. Carnegie Mus.
17 (1): 167-188, pi. 8.

Ortmann, A. E. and B. Walker. 1922. On the
nomenclature of certain North American na-
iades. Occ. Papers, Mus. Zool., Univ. Michi-
gan, no. 112, pp. 1-75.

Parmalee, p. W. 1965. A comparison of past
and present populations of fresh-water mus-
sels in southern Illinois. Trans. Illinois State
Acad. Sci. 49: 184-192.

. 1967. The fresh-water mussels of Il-
linois. Illinois State Mus. Popular Sci. Ser.
8, pp. i-ix, 1-108, 35 pis.

RoHERTSON, I. C. S. AND C. L. Blakeslee. 1948.
The Mollusca of the Niagara frontier region.
Bull. Buffalo Soc. Nat. Sci. 19 (3): i-.xi,
1-191, 14 pis., 1 folding map.

RosEWATER, J. 1959. Mollusks of the Salt River,
Kentucky. Nautilus 73: 57-63.

Say, T. 1830—1834. American Conchology, or de-
scriptions of the shells of North America, il-
lustrated by colored figures. New Harmony,
Indiana. 68 col. pis., with letterpress, pts.
1-7.

ScAMMOx, R. E. 1906. The Unionidae of Kan-
sas. Kansas Univ. Sci. Bull. 3(9): 279-372,
pis. 62-85.

Shoup, C. S. 1974. A bibliography of the zo-
ology of Tennessee and the Tennessee Valley
region. Mollusca, pp. 38-62. 251 p. U. S.
Atomic Energy Comm. Office of Info. Serv.,
Tech. Info. Office, Oak Ridge, Tennessee.

Simpson, C. T. 1900a. Synopsis of the naiades,
or pearly fresh-water mussels. Proc. U. S.
Natl. Mus. 22: 501-1044, pi. 18.

. 1900b. The evidence of the Unionidae

regarding the foniier courses of the Tennes-
see and other southern rivers. Science, n. s.
12: 133-136.

. 1914. A descriptive catalogue of the

naiades, or pearly fresh-water mussels. De-
troit, Michigan. Parts 1-3, pp. i-xi, 1-1540.

Stansberv, D. H. 1961. The naiades (Mol-
lusca, Pelecypoda, Unionidae ) of Fishery
Bay, South Bass Island, Lake Erie. Sterkiana,
no. 5: 1-35, 5 pis.

. 1964. The Mussel (Muscle) Shoals of

the Tennessee River revisited. Amer. Mai.
Union, Repts. for 1964, pp. 24-26.

. 1965. The naiad fauna of the Green

River at Munfordville, Kentucky. Amer. Mai.
Union, Repts. for 1965, pp. 13-14.

. 1970. American Malacological Union

Symposium: rare and endangered mollusks.
( 2 ) . Eastern freshwater mollusks, ( 1 ) . The
Mississippi and St. Lawrence River systems.
Malacologia 10: 9-31, pis. 1-2.

. 1971. Rare and endangered mollusks in

eastern United States, pp. 5-18. [In] S. E.
Jorgensen and R. Sharp (Editors) Proc. of
a symposium on rare and endangered mol-
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Wild., Dept. Interior, Region 3, Twin Cities,
Minn., pp. 1-79.

. 1973. A preliminary report on the na-
iad fauna of the Clinch River in the southern
Appalachian mountains of Virginia and Ten-
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Bull. Amer. Mai. Union for 1972: 20-22.

. 1976. Naiad mollusks [in] Endangered

and threatened plants and animals of Ala-
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42-52.
Stansbery, D. H. and W. J. Clench. 1975.
The Pleuroceridae and Unionidae of the Mid-
dle Fork Holston River in Virginia. Bull.
Amer. Mai. Union for 1974: 51-54.

290 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Starrett, W. C. 1971. A survey of the mus-
sels (Unionacea) of the llHnois River: a pol-
hited stream. Illinois Nat. Hist. Sur., Bull. 30
( 5 ) : 267-403, 4 col. pis., 17 figs.

Stein, F. 1881. Synopsis of the molluscous
fauna of Indiana. 16th Ann. Rept. Indiana
Gcol. Survey, pp. 451-467.

Thiele, J. 1934. Handbuch der systematischen
W'eichtierkunde, G. Fischer, Jena. 2: I-V,
779-1154.

Utterback, W. I. 1916. The naiades of Mis-
souri. Amer. Mid. Nat. 10: 432-464, pis. 27-
29. The entire work (1915-16) was repaged
and issued as a separate volume; the cited
pages were renumbered: 169-200.

VAN der Schalie, H. 1936. The naiad fauna of
the St. Joseph River drainage in southwestern
Michigan. Amer. Mid. Nat. 17: 523-527.

. 1938a. The naiad fauna of the Huron

River, in south-eastern Michigan. Misc. Pub.
Mus. Zool, Univ. Michigan, no. 40, pp. 83,
12 pis., 1 folding map.

. 1938b. The naiades (freshwater mus-
sels) of the Cahaba river in northern Ala-
bama. Occ. Papers, Mus. Zool., Univ. Michi-
gan, no. 392, pp. 1-29.

. 1939a. Mcdioiiidiis mcgJameriae, a

new naiad from the Tombigbee River, with
notes on other naiads of that drainage. Occ.
Papers, Mus. Zool., Univ. Michigan, no. 407,
pp. 1-6, 1 pi.

. 1939b. Additional notes on the naiads

( fresh-water mussels ) of the lower Tennessee
River. Amer. Mid. Nat. 22: 452-457.

. 1939c. Distributional studies of the na-
iades as related to geomorphology. Jour.
Geomorph. 3: 251-257.

— . 1941a. Zoogeography of naiades in the
Grand and Muskegon rivers of Michigan as
related to glacial history. Papers Michigan
Acad. Sci., Arts and Letters 26: 297-310.

. 1941b. The taxonomy of naiades inliabit-

ing a lake environment. Jour, of Conch. 21
(8): 246-253.

. 1945. The value of mussel distribution

in tracing steam confluence. Papers of the
Michigan Acad. Sci. Arts and Letters for
1944, 355-373.

. 1961. The naiad (fresh-water mussel)

fauna of the Great Lakes. Great Lakes Res.
Div., Inst. Sci. and Tech., Univ. Michigan,
Pub. no. 7: 156-157.

. 1973. The mollusks of the Duck River

drainage in central Tennessee. Sterkiana, no.
52, pp. 45-55.

Walker, B. 1910. Notes on TninciUa, with a
key to the species. Nautilus 24: 75-81.

. 1918. A synopsis of the classification

of the fresh-water Mollusca of North America,
North of Mexico, and a catalogue of the more
recently described species, with notes. Mus.
Zool., Univ. Michigan. Misc. Pub. no. 6, pp.
1-213.

Wilson, C. B. and H. W. Clark. 1912a. Mus-
sel beds of the Cumberland River in 1911.
Econ. Circ. no. 1. U. S. Bur. Fisheries, Wash-
ington, D. C, pp. 1-8.

. 1912b. The mussel fauna of the Maumee

River. U. S. Bur. Fisheries Doc. no. 757,
pp. 1-72, 2 pis.

. 1914. The mussels of the Cumberland

River and its tributaries. U. S. Bin-. Fisheries
Doc. no. 781, pp. 1-67, pi. 1.

Plagiola from Middle North America • Johnson 291

INDEX

an^uluta Rafinesqiie, Amhlcnia tonilosa 261

arcacfunnis (Lea), Pkigiohi (Plagiola) 257

arcaeformis Lea, Unio 257

hicnuirginata (Lea), Plagiola (Tonilosa) 268

bicmarginatus Lea, Unio 268

biloba Rafinesque, Epioblasma 282, 283

hrcviclcns Lea, C/»/o 252

capillaris Lea, (7n!0 276

Capsaefortnis Frierson 260

capsacfonnis (Lea), Plagiola (Tonilosa) 269

capsaefonnis Lea, Unio 269

cincinnatiensis Lea, L//i/o 261

c/flt'fl (Lamarck), Plcurobema 279

compactus Lea, L/ju'o 255

Crenodonta Schluter 248

cuneaiiis Swainson, f/n/o 249

ciirtisii Frierson and Utterback, TnmciUa 272

Cyprogcnia Rafinesque 243

ddicata Simpson, Tnincilla (Scalenaria)

sulcata 261
deviatus Reeve, Unio 274
Dysnomia Agassiz 282
FJlipsaria Rafinesque 248
ellipsaria Rafinesque, Obliquaiia 248
Epiohlasma Rafinesque 282
flagellatus Say, Unio 278
flexuosa R;ifinesque, Obliquaiia 283
flcxuosa (Rafinesque), Plagiola (Epioblasma) 283
floientina (Lea), Plagiola (Tondosa) 271
florentinus Lea, (7/i/o 271
foliatus Hildreth, Unio 283
fornwsus Lea, C7»/'o 249
gibbosa Rafinesque, Amblema 261
gubernaculum Reeve, (7»/o 261
haijsiana (Lea), Plagiola (Pilia) 280
hatjsiana Lea, C/»/o 280

interrupta Rafinesque, Obliquaiia (Plagiola) 252
interrupta ( Rafinesque ) Plagiola ( Plagiola ) 252
lefeveri Utterback, Tnincilla 274
/en/or (hea), Plagiola (Plagiola) 259
lenior Lea, (7n/o 257
/en/s Lea, f7»/o 259
lewisi Walker, Tnincilla 283
longiusculus Gregorio, L/zn'o triangularis 249
mcnkiana (Lea), Lampsilis 254
metastriatus Conrad, C^in'o 254
modicclltts Lea, L^»/o 255
monodonta Say, Cumheilandia 243
nebidosa (Conrad), Villosa 260
ne.r».s' Say, L^/u'o 257

»!/.v Kiister, L^»/o 274

Obliquaria Rafinesque 248

Obliqiiata Frierson 276

obliquata Rafinesque, Obliquaria 278

obliquata (Rafinesque), Plagiola (Pilea) 278

obliquus Potiez and Michaud, Unio 261

othcaloogenis Lea, Unio 254

pectitis Conrad, U;iio 278

Penita Frierson 247

penita (Conrad), Plagiola (Plagiola) 254

penitus Conrad, Unio 254

pergibosus Gregorio, Unio triangularis 249

perohliquus Conrad, Unio gibbosus 261, 278

perplcxus Lea, Unio 261

perplexus Say, [/n/o 278

personata (Say), Plagiola (Pilea) 276

personatus Say, (7»/o 276

phiUipsii Reeve, C/n/o 261

Pdea Simpson 276

pileus Lea, Unio 276

Plagiola Rafinesque 247, 252

Plagiolopsis Thiele 248

propestdcatus Gregorio, Unio 278

propinqua (Lea), Plagiola (Tondosa) 266

propinquus Lea, Unio 266

pijramidatus (Lea), Pleuiobcma 279

rangianus Lea, C/n/o 261

ridibundus Say, L/j«'o 278

saccatus Krister, L^»/o 271

sacculus Reeve, Unio 271

sampsoni Lea, Plagiola (Torulosa) 265

sampsonii Lea, (7n/o 265

Scalenilla Ortmann and Walker 276

sowerbyanus Lea, t/jiio 280

steicardsoni (Lea), Plagiola (Epioblasma) 285

steivardsoni Lea, Un/o 285

ifeii'ejisoJH' Gregorio, C7(h'o stewardsonii 278

stdcatus Lea, C7ji/o 278

Torulosa Frierson 260

torulosa Rafinesque, Amblema 261

torulosa (Rafinesque), Plagiola (Torulosa) 261

triangularis Barnes, l7n/o 249

triquetra (Rafinesque), Plagiola

(TruncillopsLs) 248
triquetra Rafinesque, Tnincilla 248
Tnincillopsis Ortmann and Walker 248
turgidula (Lea), Plagiola (Torulosa) 274
turgidulus Lea, Unio 274
icalkcri Wilson and Clark, Truucilla 271

292 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 1.
The distribution of Plagiola {Truncillopsis) triquetra (Rafinesque).

This, the most primitive species of Plagiola. is the most widely distributed in the genus. It appears to have
had refugia west of the f^ississippi River beyond maximum Pleistocene glaciation (roughly to the Missouri
River) on the Old Prairie and in the Meramec Basin, as well as in the upper White River system on the Ozark
Plateau south of the Ozark Crest (record enclosed by large circle).

The occurrence of this species in the White River on the Ozark Plateau south of the Ozark Crest, as well as in
the Tennessee and Cumberland river systems, including the Duck River drainage of the former, on the Cum-
berland Plateau, suggests that this species has persisted at least since the Cretaceous uplift. P. triquetra
may also have had refugia in the Allegheny and Monongahela river drainages in the mountainous region of
western Pennsylvania and West Virginia during the Pleistocene.

A. In post-glacial time triquetra spread into Wisconsin from a connection between the Fox and Wisconsin riv-
ers in the vicinity of Portage, Columbia Co., Wisconsin.

B. The presence of triquetra in the Illinois River, Illinois and the Muskegon, Grand and St. Joseph rivers on
the eastern side of Lake Michigan, suggests that before the formation of Lake Michigan the latter streams
were tributaries of the Des Plaines River, which by way of the Chicago outlet drained into the Mississippi
River, by way of the Illinois River.

C. The distribution of triquetra in the present rivers flowing into western and southern Lake Erie indicates
they were part of the Greater Maumee River system, when the bed of Lake Erie was partially dry during the
Trent outlet stage of the Great Lakes, and that there was a connection between the Wabash and Maumee
rivers in the vicinity of Fort Wayne, Allen Co., Indiana.

Plagiola from Middle North America • Johnson 293

294 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 2.
The distribution of Plagiola (Torulosa) turgidula (Lea) (triangles), and Plagiola (Torulosa) florentina (Lea) (dots).

The restricted distribution of P. turgidula and florentina to the upper White River system on the Ozark Plateau
(records south of the Ozark Crest enclosed by large circles) and to the Tennessee and Cumberland river sys-
tems (including the Duck River drainage of the former) on the Cumberland Plateau, suggests that these species
have persisted at least since the Cretaceous uplift.

Plagiola from Middle North America • Johnson 295

296 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 3.
The distribution of Plagiola (Torulosa) iorulosa (Rafinesque).

This species has persisted in the Tennessee River system, and while it may have had refugia in the Green, Al-
legheny and Monongahela river drainages, it is assumed that it spread from the Tennessee River system in
post-glacial time to form its present distribution.

Although not now found in any intervening rivers, it must have followed the same route as triquetra (Plate 1
B) in reaching the Grand River, Michigan.

Clearly it followed the same route as triquetra (Plate 1 C) in reaching lakes Huron and Erie.

Plagiola from Middle North America • Johnson 297

298 Bulletin Museum of Comporotivc Zoology, Vol. 148, No. 6

Plate 4.
The distribution of Plagiola (Pilea) obliquata (Rafinesque).

This species has persisted in the Tennessee and Cumberland river systems, and while it may have had a re-
fugia in the Green River drainage, it is assumed that it spread from the former systems in post-glacial time
to form its present distribution.

Clearly it followed the same route as triquetra (Plate 1 C) and torulosa (Plate 3) in reaching Lake Erie. Un-
like the latter two species, which occur in the Allegheny and Monongahela river drainages, it is not known
to have ascended the Ohio River beyond Cincinnati, Hamilton Co., Ohio, though like P. flexuosa (Plate 5), it
probably did.

Plagiola from Middle North America • Johnson 299

300 BuUetiti Museum of Comparative Zoology, Vol. 148, No. 6

Plate 5.
The distribution of Plagiola (Epioblasma) flexuosa (Rafinesque).

This species has persisted in the Tennessee and Cumberland river systems; it is assumed that it spread from
one or both of these systems in post-glacial time into the Ohio River system, where it occurs from the lower
Wabash River drainage to the Ohio River, Steubenville, Jefferson Co., Ohio.

Pl.AGlOLA FROM MlDDLE NoRTH AmERICA • JokuSOn 301

mi.

302 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 6.

The distribution of Plagiola (Pilea) personata (Say), Fig. A, Plagiola (Torulosa) propinqua (Lea), Fig. B (dots),
and Plagiola (Torulosa) sampsoni (Lea), Fig. B (triangles).

P. personata and propinqua have persisted in the Tennessee and Cumberland river systems, w^hereas sampsoni
appears to be absent in the Cumberland. It is assumed that sampsoni spread from the Tennessee, while per-
sonata and propinqua spread from either or both the Tennessee and Cumberland river systems in post-glacial
time, into the Ohio River system. They all occur in the lower Wabash River drainage, and extend in the
Oh^o drainage to Cincinnati, Hamilton Co., Ohio.

The location of sampsoni in the Tennessee River system is conjectural. An additional record, from the Ohio
River at Cincinnati, is missing from the map.

Plagiola from Middle North America • Johnson 303

304 BuUetin Museum of Comparatwc Zoology, Vol. 148, No. 6

Plate 7.

The distribution of Plagiola (Plagiola) interrupta (Rafinesque), Fig. A, Plagiola (Torulosa) capsaeformis (Lea),
Fig. B. and Plagiola (Plagiola) lenior (Lea), Fig. C.

These species have persisted in the Tennessee and Cumberland river systems, and the Duck River drainage
of the former. According to Ortmann (1924: 46), Ducl< River v^^as originally more directly connected with the
Tennessee and Cumberland rivers.

They did not extend their distribution in post-glacial time.

Plagiola from Middle North America • Johnson 305

/-:>:::^^&^f

— -■■,- V

CM^''

lo.l'

ly.

;^.

i>.i.

Hi' Win.vll

.UK*

rU-^l!l^■

vv.

/ \

i:*t

ir. .r.

,,:l-t ■"'■"-M

•,) i-

■■^-fl

-TV"^/-/ /\-M.n.ci> r\'^V'^\-^AH ^--v- { '-

306 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 8.

The distribution of Plagiola (Pilea) hayslana (Lea), Fig. A, Plagiola {Plagiola) arcaeformis (Lea), Fig. B,
and Plagiola (Epioblasma) stewardsoni (Lea), Fig. C.

These species have persisted in the upper and lower Tennessee River system as w/ell as in the Cumberland
River system. They did not extend their distribution in post-glacial time.

Plagiola from Middle North America • Johnson 307

308 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 9.

The distribution of Plagiola (Torulosa) biemarginata (Lea), Fig. A, and Plagiola (Plagiola) penita (Conrad),
Fig. B.

Figure A. P. (7.) biemarginata (Lea) has persisted in the upper and lower Tennessee River system and
the Big South Fork of the Cumberland River. It, like the species shown on Plate 8, did not extend its dis-
tribution in post-glacial time.

Figure B. P. (P.) penita Conrad is restricted to the Alabama-Coosa River system and is clearly derived from
P. (P.) interrupta (Rafinesque) of the Tennessee River system.

Plagiola from Middle North America • Johnson 309

310 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 10.
Plagiola (Truncillopsis) triquetra (Rafinesque)

Figure 1. Truncilla triqueter Rafinesque. Falls of the Ohio River, [near Louisville, Jefferson Co., Kentucky].
Lectotype ANSP 20231. Length 55, height 37, width 25 mm. Male.

Figure 2. Green River, 8 mi. S Campbellsville, Taylor Co., Kentucky. MCZ 220157. Length 44, height 31,
width 24 mm. Male.

Figure 3. Ohio River, Cincinnati, Hamilton Co., Ohio. MCZ 6158. Length 39, height 24, width 24 mm. Fe-
male.

Figure 4. Green River, 8 mi. S Campbellsville, Taylor Co., Kentucky. MCZ 220157. Length 36, height 24,
width 22 mm. Female.

Plagiola (Plagiola) interrupta (Rafinesque)

Figure 5. Obliquaria (Plagiola) interrupta Rafinesque. [Cumberland River, Tennessee]. Lectotype ANSP
20257. Length 55.5, height 43, width 26.5 mm. Male.

Figure 6. Unio brevidens Lea. [Cumberland River, Tennessee]. Specimen subsequently identified by Lea
USNM 85349. Length 54, height 38, width 23.5 mm. Male.

Figure 7. Unio brevidens Lea. [Cumberland River, Tennessee]. Specimen subsequently identified by Lea
USNM 85349. Length 68, height 53.5, width 43.5 mm. Female.

Plagiola (Plagiola) penita (Conrad)

Figure 8. Coosa River, Weduska Shoals, Shelby Co., Alabama. MCZ 29817. Length 52, height 40, width 26
mm. Male.

Figure 9. Unio penitus Conrad. Alabama River, near Claiborne, Munroe Co., Alabama. Holotype ANSP
59860. Length 51, height 35, width 26 mm. Female.

Figure 10. Unio metastriatus Conrad. Black Warrior River, near Blount's Springs, Blount Co., Alabama.
Length 30, height 23, width 17 mm. Syntype [lost], from Conrad. Male.

F.gure 11. Unio metastriatus Conrad. Black Warrior River, near Blount's Springs, Blount Co., Alabama.
Length 28, height 21 mm, width unavailable. Syntype [lost], from Conrad. Female.

Figure 12. Unio compactus Lea. Etowah River, Georgia. Holotype USNM 84447. Length 32, height 27, width
19 mm. Male.

Figure 13. Unio compactus Lea. Etowah River, Georgia. Allotype USNM 84447a. Length 28, height 20, width
16 mm. Female.

Figure 14. Unio modicellus Lea. Connasauga River, Georgia. Holotype USNM 84841. Length 28, height 21,
width 15 mm. Male.

Figure 15. Unio othcaloogensis Lea. Oothkalooga Creek, Gordon Co., Georgia. Holotype USNM 84615. Length
22, height 16, width 13 mm. Male.

Plagiola from Middle North America • Johnson 311

312 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 11.
Plagiola (Plagiola) arcaeformis (Lea).

Figure 1. Unio arcaeformis Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Specimen subsequently
identified by Lea USNM 84422. Length 54.5, height 43.5, width 38 mm. Male.

Figure 2. Cumberland River, Tennessee. MCZ 5033. Length 42, height 34, width 27 mm. Male.

Figure 3. Cumberland River, Tennessee. MCZ 5033. Length 55, height 40, width 40 mm. Female.

Figure 4. Unio arcaeformis Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Specimen subsequently
identified by Lea USNM 84422. Length 56.5, height 39, width 39 mm. Female.

Plagiola (Plagiola) lenior (Lea)

Figure 5. Eastern Tennessee. MCZ 16387. Length 35, height 21, width 14 mm. Male.

Figure 6. Unio lenior Lea. Stones River, Tennessee. Holotype USNM 86130. Length 25, height 16, width 12.5
mm. Female.

Plagiola (Torulosa) torulosa (Rafinesque)

Figure 7. Amblema gibbosa Rafinesque. Ohio River. Lectotype ANSP 20232. Length 40, height 33, width 25
mm. Male.

Figure 8. Unio perplexus Lea. Ohio River, Cincinnati, Hamilton Co., Ohio. Holotype USNM 84324. Length
64, height 43, width 32.5 mm. Male.

Figure 9. Unio cincinnatiensis Lea. Ohio River, Cincinnati, Hamilton Co., Ohio. Holotype USNM 84199. Length
48, height 39, width 27.5 mm, Male.

Figure 10. Unio rangianus Lea. Mahoning River, near Poland, Mahoning Co., Ohio. Allotype USNM 84798.
Length 51,5, height 36, width 26 mm, Male.

Figure 11. Truncilla sulcata delicata Simpson. Detroit River. Amherstburg, Essex Co., Ontario. Holotype
USNM 160853. Length 44, height 31, width 24 mm. Male.

Plagiola from Middle North America • Johnson 313

314 Btilletm Museum of Comparative Zoology, Vol. 148, No. 6

Plate 12.
Plagiola (Torulosa) torulosa (Rafinesque)

Figure 1. Amblema torulosa Rafinesque. Kentucky River, Kentucky. Holotype ANSP 20218. Length 65, height
48, width 33.5 mm. Female.

Figure 2. Unio rangianus Lea. Mahoning River, near Poland, Mahoning Co., Ohio. Lectotype USNM 84798.
Length 49, height 35.5, width 24 mm. Female. [Slightly larger than Lea's figured type].

Figure 3. Unio gibbosus perobliquus Conrad. Wabash River, Indiana. Figured type [lost] from Conrad. Length
56, height 48, width 32 mm. Male.

Figure 4. Unio phillipsii Reeve. North America. Length 54, height 45, width 30 mm. Holotype BMNH 196481.
Female.

Figure 5. Unio gubernaculum Reeve. (Hab?). Holotype BMNH 1965203. Length 57, height 48, width 19 mm.
Female.

Plagiola (Torulosa) sampsoni (Lea)

Figure 6. Unio sampsonii Lea. Wabash River, New Harmony, Posey Co., Indiana. Holotype USNM 84802.
Length 42.5, height 33, width 25.5 mm. Male.

Figure 7. Unio sampsonii Lea. Wabash River, New Harmony, Posey Co., Indiana. Allotype USNM 84802.
Length 45, height 38.5, width 29 mm. Female.

Plagiola (Torulosa) propinqua (Lea)

Figure 8. Unio propinquus Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Holotype USNM 84332.
Length 53, height 41, width 33 mm. Male.

Figure 9. Holston River, Knoxville, Knox Co., Tennessee. MCZ 5819. Length 44, height 46, width 31 mm.
Female.

Plagiola from Middle North America • Johnson 315

316 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 13.
Plagiola (Torulosa) biemarginata (Lea)

Figure 1. Unio biemarginatus Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Allotype USNM
84608a. Length 38.5, height 30, width 19 mm. Male.

Figure 2. Unio biemarginatus Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Holotype USNM
84608. Length 36, height 31.5, width 18 mm. Female.

Plagiola {Torulosa) capsaeformis (Lea)

Figure 3. Unio capsaelormis Lea. Cumberland River, Tennessee. Holotype MCZ 178570. Length 45, height
33, width 24 mm. Male.

Figure 4. Unio capsaeformis Lea. Cumberland River, Tennessee. Specimen subsequently identified by Lea
MCZ 178568. Length 43, height 30, width 18 mm. Female.

Plagiola (Torulosa) florentina (Lea)

Figure 5. Unio florentinus Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Allotype USNM 84948.
Length 36, width 27, height 20 mm. Male.

Figure 6. Truncilla walkeri Wilson and Clark. East Fork of Stones River, near Walterhill, Rutherford Co., Ten-
nessee. Paralectotype MZUM 90729. Length 47, height 34, width 22 mm. Male.

Figure 7. Truncilla curtisii Frierson and Utterback. White River, Forsyth, Taney Co., Missouri. Paralectotype
MZUM 90748. Length 35, height 26, width 19 mm. Male.

Figure 8. Unio florentinus Lea. Tennessee River, Florence, Lauderdale Co., Alabama. Holotype USNM
84948. Length 33, height 27, width 19 mm. Female.

Figure 9. Truncilla walkeri Wilson and Clark. East Fork of Stones River, Walterhill, Rutherford Co., Tennes-
see. Lectotype MZUM 90729. Length 42, height 30, width 17 mm. Female.

Figure 10. Truncilla curtisii Frierson and Utterback. White River, Forsyth, Taney Co., Missouri. Lectotype
MZUM 90748. Length 32, height 27, width 17 mm. Female.

Figure 11. Black River, Hendrickson, Butler Co., Missouri. MCZ 260979. Length 26, height 21, width 13
mm. Female.

Figure 12. Unio sacculus Anthony. Tennessee. Holotype MCZ 161898. Length 33, height 24, width 16 mm.
Female.

Figure 13. Truncilla curtisii Frierson and Utterback. White River, Hollister, Taney Co., Missouri. Figured
syntype [lost] from Utterback. Length 33, height 23, width 15 mm. Female.

Figure 14. Truncilla curtisii Frierson and Utterback. White River, Hollister, Taney Co., Missouri. Figured
syntype [lost] from Utterback. Length 22.5, height 19.5, width 13.5 mm. Male.

Plagiola from Middle North America • Johnson 317

318 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 14.
Plagiola (Torulosa) turgidula (Lea)

Figure 1. Truncilla lefevrei Utterback. Black River, Williamsvllle, Wayne Co., Missouri. Figured syntype [lost]
from Utterback. Length 32, height 21, width 15 mnn. Female.

Figure 2. Truncilla lefeveri Utterback. Black River, Williamsville, Wayne Co., Missouri. Figured syntype [lost]
from Utterback. Length 26.5, height 18, width 14 mm. Male.

Figure 3. Unio turgidulus Lea. Cumberland River, Tennessee. Holotype USNM 84946. Length 41.5, height
30, width 23.5 mm. Male.

Figure 4. Spring Creek, Hardy, Sharp Co., Arkansas. MZUM 90742. Length 40, height 29, width 17 mm. Male.

Figure 5. Unio deviatus Anthony. Tennessee. Holotype MCZ 161895. Length 39, height 25, width 19 mm.
Female.

Figure 6. Spring Creek, Hardy, Sharp Co., Arkansas. MZUM 90742. Length 33, height 24, width 15 mm. Fe-
male.

Plagiola (Pilea) personata (Say)

Figure 7. Unio pileus Lea. Ohio River, near Cincinnati, Hamilton Co., Ohio. Holotype USNM 84602a. Length
47, height 43, width 28 mm. Male.

Figure 8. Unio capillaris Lea. Ohio. Specimen subsequently identified by Lea USNM 84602. Length 54,
height 46, width 33 mm. Female.

Figure 9. Unio personatus Say. Cumberland River, Tennessee. Neotype MCZ 5763. Length 54, height 48,
width 31 mm. Female.

Plagiola (Pilea) obliquata (Rafinesque)

Figure 10. Obliquaria obliquata Rafinesque. Kentucky River. Lectotype ANSP 20226. Length 59, height 43,
width 32.5 mm. Male.

Figure 11. Unio sulcatus Lea. Ohio. Holotype USNM 84803. Length 57, height 41, width 33 mm. Male.

Figure 12. Unio sulcatus Lea. Ohio. Allotype USNM 84803. Length 37, height 29.5, width 23 mm. Female.

Plagiola from Middle North America • Johnson 319

320 Bulletin Museum of Comparative Zoology, Vol. 148, No. 6

Plate 15.
Plagiola (Pilea) haysiana (Lea)

Figure 1. Unio haysianus Lea. Cumberland River, Nashville, Davidson Co., Tennessee. Specimen subse-
quently identified by Lea USNM 84613. Length 51.5, height 49, width 36 mm. Male.

Figure 2. Unio sowerbyanus Lea. Caney Fork of the Cumberland River, Tennessee. Specimen subsequent-
ly identified by Lea MCZ 178886. Length 39, height 38, width 28 mm. Male.

Figure 3. Cumberland River, Tennessee. MCZ 5451. Length 32, height 30, width 28 mm. Female.

Figure 4. Unio haysianus Lea. Cumberland River, Nashville, Davidson Co., Tennessee. Specimen subse-
quently identified by Lea USNM 84613. Length 25.5, height 24.5, width 19 mm. Female.

Plagiola (Epioblasma) flexuosa (Rafinesque)

Figure 5. Obliquaria flexuosa Rafinesque. Kentucky River. Lectotype ANSP 20249. Length 57, height 47,
width 33 mm. Male.

Figure 6. Truncilla lewisi Walker. Holston River, Tennessee. Paralectotype MZUM 91456. Length 43,
height 37, width 22.5 mm. Male.

Figure 7. Epioblasma biloba Rafinesque. Ohio River, Cincinnati, Hamilton Co., Ohio. Neotype ANSP 56571.
Length 65, height 59, width 40 mm. Female.

Figure 8. Truncilla lewisi Walker. Holston River, Tennessee. Lectotype MZUM 91456. Length 51, height
49.5, width 25 mm. Female.

Plagiola (Epioblasma) stewardsoni (Lea)

Figure 9. Unio stewardsoni Lea. Tennessee. Specimen subsequently identified by Lea MCZ 178817. Length
31, height 29, width 18 mm. Male.

Figure 10. Unio stewardsoni Lea. Tennessee River, Chattanooga [Hamilton Co.], Tennessee. Holotype
ANSP 56572. Length 31, height 29, width 13 mm. Female.

Plagiola from Middle North America • Johnson 321

ISSN 0027-4100

SuliQtln OF THE

Museum of

Comparative

Zoology

Biology and Evolution of

the Avian

snus

Atlapetes (Emberizinae

RAYMOND A. PAYNTER, JR.

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 7
5 JULY 1978

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SPECIAL PUBLICATIONS.

1. VVhittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and
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2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini-
dae (Mollusca: Bivalvia). 265pp.

3. Sprinkle, J., 1973. Moi-phology and Evolution of Blastozoan Echinoderms.
284 pp.

4. Eaton, R. J. E., 1974. A Flora of Concord. 236 pp.

Other Publications.

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Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa-
chusetts, 02138, U.S.A.

BIOLOGY AND EVOLUTION OF THE AVIAN GENUS
ATLAPETES (EMBERIZINAE)*

RAYMOND A. PAYNTER, JR.^

CONTENTS

Abstract 323

[ntroduction 324

Acknowledgments 326

Distribution, Habits, and Morphology 327

Atlapetes alhinucha superspecies 327

Atlapetes albinucJui 327

Atlapetes pallidinucha 330

Atlapetes nifiniicha superspecies 330

Atlapetes riifiniicha 330

Atlapetes melanocephalus 334

Atlapetes tricolor superspecies 334

Atlapetes flaviceps _ 334

Atlapetes fuscoolivaceus 334

Atlapetes tricolor .___ 335

Atlapetes albofrenatus 336

Atlapetes leucopis 337

Atlapetes pileatus 338

Atlapetes fulviceps superspecies 339

Atlapetes semirufus 339

Atlapetes personatus ____ 340

Atlapetes fulviceps 341

Atlapetes citrinellus 341

Atlapetes brunneinucha __. 341

Atlapetes torquatiis superspecies 345

Atlapetes virenticeps 347

Atlapetes atricapillus 350

Atlapetes torquatiis 350

Origin and Interrelationships of the Taxa 353

Introduction 353

^ Museum of Comparative Zoology.
* Published and supported in part by a grant
from the Wetmore Coles Fund.

Atlapetes albinucha superspecies 354

Atlapetes rufinuclia species-group 356

Atlapetes rufinuclia superspecies 356

Atlapetes tricolor superspecies 357

Atlapetes albofrenatus 359

Atlapetes leucopis - 361

Atlapetes pileatus 361

Atlapetes fulviceps superspecies 361

Atlapetes citrinellus 362

Atlapetes schistaceus species-group 362

Atlapetes schistaceus 362

Atlapetes nationi 362

Atlapetes leucoptenis 364

Atlapetes albiccps 364

Atlapetes pallidiceps 364

Atlapetes nifigenis - — - 364

Atlapetes torquatus species-group 364

Atlapetes brunneinucha — 364

Atlapetes torquatus superspecies 365

Conclusions 366

Literature Cited 368

Abstract. The 24 species of Atlapetes are
furtive brush or forest inliabitors occurring mainly
in subtropical or temperate zones in the higlilands
from Mexico to the southern Andes. The\- are
grouped into four subunits: (1) A. albinucha
superspecies, (2) A. rufinuclia species-group with
A. rufinuclia superspecies, A. tricolor superspecies,
A. albofrenatus, A. leucopis, A. pileatus, A. fulvi-
ceps superspecies, and probably A. citrinellus, (3)
A. schistaceus species-group with A. schistaceus,
A. nationi, A. Icucopterus, A. albiceps, A. pallidi-
ceps, and A. rufigenis, and (4) A. torquatus spe-
cies-group with A. brunneinucha and A. torquatus
superspecies. The A. rufinuclia species-group,
with 11 (or 12) species, and the A. schistaceus
species-group, with six species, are the main evolu-

Bull. Mus. Comp. ZooL, 148(7): 323-369, July, 1978 323

324 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

tionary lines. Their re.specti\e members are largel>-
allopatric, suggesting a recent origin. Speciation
witiiin Atlapctcs seems to ha\e been greatly in-
fluenced by Pleistocene climatic ocellations and to
ha\e been most acti\'e in the northern Andes.

INTRODUCTION

Members oi the genu.s AtJapetes, a moder-
ately well-differentiated genus of emberi-
zine sparrows, occur from tlie plateau of
northern Mexico through the mountains of
Central America and down the length of
the Andes to northwestern Argentina (Fig.
1). There are 24 species, with the greatest
number occurring in Colombia, where there
are 14 species, and in Ecuador, where there
are 11 species (Fig. 2). There is a general
attenuation in the number of species north
and south of the Colombia-Ecuador region.
Several species have ranges that are re-
stricted to a single valley system (flaviceps;
palUdiceps) or massif {melanocephalus) ,
while at the other extreme one species
( brtinnei nucha) occurs from Mexico to Pern
and has "a more extended range, . . . , than
that of any other subtropical . . ." bird
(Chapman, 1923a: 245). However, for the
most part, species of Atkipetes have ranges
that are intermediate in length. The distri-
bution of a number of species is discontin-
uous; at times the breaks appear to be
unrelated to geography, vegetation, or col-
lecting activities and offer clues to the
evolutionary history of the genus.

All species are roughly the same size ( ca.
130-180 mm long) and generally rather
plainly marked. They are brown, brownish
gray, or dark green dorsally and dull gray
or bright yellow below. The majority have
brown heads, the others are black with the
exception of one which is white.

Members of the genus are character-
istically furtive, seldom ascending more
than a meter or two above the ground or
emerging from the dense vegetation they

prefer. They are solitary during the breed-
ing season but may assemble in what appear
to be family groups at other seasons. While
many brush-inhabitors have loud, easily
recognized vocaUzations, the calls and
songs of members of the genus Atkipetes are
faint and undistinctive. In some localities I
these birds seem to make up a substantial
portion of the avifauna, but this is apparent
only to the most diligent observer.

Considerable information is available on
the distribution of the genus. This has pro-
vided material for my primary objective,
which is to reconstruct the evolutionary
history of the species of Atkipetes. The sec-
ondary purpose of this paper is to assemble
what information is available on all aspects
of this little-known group. Knowledge of
the living birds is so scant that the best
that can be done is to establish a foundation
for future studies. i

The arrangement of species in Peters'
Check-list (Paynter, 1970:190-206) was
based on a preliminary study of the genus.
The present treatment is essentially the
same, except that two taxa ( virenticeps and
atricapiUus) that were treated as races of
A. torqiiatus are now considered to be al-
k)species of the torcjuatiis superspecies. I
have also somewhat revised the sequence of
species to reflect my present concept of
their relationship.

Information on the distribution and habits
of the genus has been gathered from pub-
lished material, from a limited amount of
my own fieldwork ( principally in Mexico,
Colombia, and Ecuador) and from the ex-
amination of museum specimens (mainly
the rarer species or those with limited
ranges). The occurrences of all 24 species
have been plotted on the accompanying
distribution maps. \\'ithout doubt some ex-
isting records have not been found, par-
ticularly among the many specimens of
the common species, but additional data
from collections should not materially alter

Figure 1. Distributional records for the genus Atlapetes.

Biology and Evolution of Atlapetes • Paynter 325

326 Bulletin Museum of Comparative ZooJogij, Vol. 148, No. 7

MEXICO

GUATEMALA

HONDURAS

EL SALVADOR

COSTA RICA

PANAMA, western

PANAMA, eastern

VENEZUELA, PANTEPUI

VENEZUELA, ANDES

COLOMBIA, SANTA MARTA

COLOMBIA, E. ANDES

COLOMBIA, C. ANDES

COLOMBIA, W. ANDES

ECUADOR

PERU, north of Maranon

PERU, south of Maranon

BOLIVIA

ARGENTINA

1 3 5 7 9 11
I I I 1

'*!•!"''.*.'•'

^^ = endemic

• ' '.

Hi::::::

■

•!\\*!'''I'!'.\vI*^"I'I"

:j:j:::x:x:v:::::::::::::v:::::v::::

|||^^^^^^^;•:•;■:•:•^■^:■;•:■:•;■;'.■.■.■.•.■.^■.

Rfe::::::::;:;:;:;:;:;:;:?w::::::¥S¥S

Figure 2. Geographical distribution of tine 24 species of Atlapetes.

the depicted ranges of the more aljiindant
forms. On the other hand, additional field-
work should reveal more extensive ranges
for at least some of the rarer species and
for some of those whose distribution is now
believed to be very restricted. Furthermore,
one should not fail to appreciate that these
maps show the distribution of each species
from the time of its discovery until now.
Therefore, the maps depict the maximum
known ranges of the species. The enormous
changes wrought on Neotropical vegeta-
tion during the past 100 years may have
made the present-day ranges of some spe-
cies more restricted than shown on the
maps, while other forms may have flour-
ished under these changes and occur out-
side the ranges as plotted.

To avoid repetition and cumbersome de-
tail, I have not documented, by means of

literature citations or reference to museum
specimens, every locality plotted on the
distribution maps or mentioned in the text.
The compilation of Hellmayr (1938:384-
423) was the primary source; records from
more recent publications and from museum
material are on file and available to futvire
workers.

The A. schistaceus species-group, con-
taining A. schistaceus, mitioni, leucopterus,
alhiceps, pallidiceps, and riifi^enis, was
analysed in an earlier paper ( Paynter, 1972)
and will not be treated in detail again.

ACKNOWLEDGMENTS

I am grateful to the following institutions
and individuals who have loaned specimens
and answered queries regarding their col-
lections: Dean Amadou of the American

Biology and Evolution of Atlapetes • Paijnter 327

Museum of Natural History; Melvin A.
Traylor, Jr. and Emmet R. Blake of the
Field Museum of Natural History; James
Bond, Frank Gill, and R. M. de Schauensee
of the Academy of Natural Sciences of
Philadelphia; John W. Hard\% formerly of
the Moore Collection at Occidental College;
George H. Lowery, Jr. of the Museum of
Zoology, Louisiana State University; Robert
W. Storer of the Museum of Zoology, Uni-
versity of Michigan; Kenneth E. Stager of
the Los Angeles County Museum; and Ernst
Sutter of the Naturhistorisches Museum
Basel.

For sharing with me their field experi-
ences with a number of species of Atlapetes
I thank Paul K. Donahue, John W. Fitz-
patrick, John Terborgh, and Richard E.
Webster.

The following have read drafts of the
manuscript and offered me their comments,
for which I thank them: J. W. Fitzpatrick,
J. Haffer, E. Mayr, M. A. Traylor, Jr., and
R. E. Webster.

Alison Pirie has prepared the maps, typed
the manuscript, and done those innumerable
but necessary chores that only the author
can appreciate.

DISTRIBUTION, HABITS, AND
MORPHOLOGY

Atlapetes albinucha superspecies

There are two allospecies in this superspe-
cies, viz. A. albinucha and A. paUidiniicha.
Morphologically, they are only moderately
similar, but it is their geographical and
altitudinal distribution that leads me to
believe that thev share a common ancestor.

Atlapetes albinuchia

Range. — A. albinucha is primarily a Mid-
dle American species of middle altitudes. It
occurs from the Caribbean slope of central
Mexico (Veracruz and Puebla), and the
Pacific slope of southwestern Mexico (Chi-
apas), through the highlands of Central
America to western Panama (Figs. 3, 4).
It reappears in Colombia, where it occurs

on all three ranges of the Andes except in
the extreme south (Nariiio) and in the
Eastern Cordillera north of Cundinamarca.

Tills is a form principally of the humid
subtropical and lower temperate zones.
It has been found as low as 600 m in Mexico
(Miller et al., 1957) and Costa Rica
(Skutch, 1967) and up to 3,050 m in Guate-
mala (Land, 1970), but seems to occur
most often from about 1,000 to 2,400 m. No
latitudinal variation in altitudinal distribu-
tion is evident, in spite of Skutch 's (1967)
impression that the species occurs at higher
elevations in southern Central America than
in Mexico and Guatemala.

Halyitat. — This atlapetes is found in thick
\'egetation near the edges of forests, in
brushy woodland, and in dense second-
growth, but not in the interior of heavy
forest. It does not seem to occur regularly
in pure stands of conifers, although Skutch
( 1967 ) once found it in low cypress. It is,
however, common in mixed pine-broadleaf
associations and in the low deciduous vege-
tation bordering pine forest ( Dickey and
^'an Rossem, 1938; Pavnter, 1957; Skutch,
1967).

Habits. — A. albinucha is secretive and
difficult to observe, as are all species of
Atlapetes. Skutch (1967) reports that this
species is even more shy than A. brun-
neinucha and A. torquatus. It generally oc-
curs on or near the ground, but at times will
go as high as 20 m (Slud, 1964); it roosts in
trees ( Skutch, 1967 ) and occasionally feeds
there (Dickey and van Rossem, 1938;
Skutch, 1967). Except when mated or ac-
companied by its young, the species is soli-
tary (Skutch, 1967), sulking in the under-
brush and seldom flying, and then only for
short distances "in fluttering dips with [its]
tail partially cocked" (Slud, 1964:282).

\Miile in Honduras in April 1976, I saw
this species for the first time in many years,
and my attention was immediately drawn
to the loud noise of the flapping wings, a
characteristic I had first noticed in A. na-
tioni (Paynter, 1972:303). One wonders
whether the sound of the wingbeats, in-

328 Bulletin Mu6ciim of Comparative Zoology, Vol. 148, No. 7

Figure 3. Northern races of A. albinucha.

stead of voice signals, is a means of keep-
ing members of a family gionp together, or
whether the noise is merely an aerodynamic
byproduct with no special significance.

Skutch (1967) saw the species feeding
on berries of Fuchsia arhorescens. This
seems to be the only specific record of its
diet but because the bird is often heard
foraging in the forest litter, it may be as-
sumed to be omnivorous. Skutch (1967)
surmises that the bird turns the litter with
its bill, although apparently no one has
yet seen this behavior.

A. albinucha builds a bulky, open nest in
weeds close to or on the ground or in dense
tangles a meter or two above it (Cherrie,
1892; Blake, 1956; Skutch, 1967). Two or
three white or pale blue eggs are laid, and
at least some white eggs turn pale blue
when their contents are removed (Cherrie,
1892). The nests are frequently parasitized

l)v Muluthrus aeneus (Cherrie, 1892; Slud,
1964).

The breeding season is poorly known. In
Middle America, it appears to be from April
through June or even late Julv (Cherrie,
1892; Blake, 1958; Paynter, 1957; Skutch,
1967); in Colombia, March and April dates
are recorded ( Miller, 1963 ) .

The call is said to be a faint high-pitched
tseep-t.see-eep (Edwards, 1972) or sst, sr, or
tsr (Slud, 1964), and its song is described
as a weak "squeaky pully" sound with
descending churrs ( Slud, 1964 ) on a thin
slow "O see me, O see, I'm weary, pity me"
(Skutch, 1967).

Morphulooical variation. — There is minor
sexual dimorphism in size, with the male's
wing and tail averaging slightly longer and
the bill minutely longer. No geographical
variation in these characters is apparent.

There is very little variation in the color

Biology and Evolution of Atlapetes • Paynter 329

Figure 4. Southern races of A. albinucha.

of the populations from southernmost Mex-
ico southward. The differences that do
exist are cHnal changes in the intensity of
the yellow of the throat ( and possibly in its
extent ) , in the degree of brownish or black-
ish cast on the back, and in the grayness of
the flanks. The variation in back color, how-
ever, may merely be a function of the age
of museum skins. From southwestern Mex-
ico to Colombia seven races have been rec-
ognized (Paynter, 1970) on the basis of
color, but these are very weak races and at
least two (colomtus and azuerensis) are al-
most certainly invalid. On the other hand,
an eighth race, the population of eastern
Mexico (nominate aJJnnucha) is conspicu-
ously different in that the entire underparts,
rather than merely the throat, are yellow.
There are no other differences.

A. a. aUnmicha was long treated as a dis-
tinct species. This is doubtless because it
was originally described from Cartagena
on the coast of Colombia ( a most improb-
able locality for any form of Atlapetes),

and, consequently it was thought to
occur only in Mexico and Colombia. Payn-
ter (1964) pointed out that the collector
of the holotype had visited both Co-
lombia and eastern Mexico on the same
voyage, and because there has never been
another specimen of the taxon collected in
Colombia, it is logical to assume the prove-
nance of the type was wrongly recorded.
The type locality was, therefore, amended
to the Caribbean slope of Mexico, and
Colombia was deleted from the range of
albinucha.

A. a. albinucha does not intergrade with
A. a. griseipectiis of southwestern Mexico,
Guatemala, and El Salvador, although the
two forms are known to occur less than 100
kilometers apart in southwestern Chiapas.
It is possible that the valley of the Rio
Crijalva ( = Rio Grande ) is sufficiently low
here to act as a barrier. The river is at
approximately 1,000 m in this region while
the bird has been taken on both sides of the
\'alley at 1,500 m or higher. It could be

330 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

argued that since the two taxa are so nearly
parapatric they should be treated as allospe-
cies. Ne\ertheless, although the entirely
yellow undeiparts of A. a. alhimicha are
conspicuously different from the gray abdo-
men and yellow throat of A. a. griseipectus
and the remaining races, this dissimilarity
almost certainly is not indicative of great
genetic difference. Furthermore, it would
appear that intergradation is pre\'ented by
a geographical barrier (low valley) and is
not the result of ecological exclusion. I find
it difficult to conceive that interbreedino;
would not occur if the taxa were in contact
and believe that racial treatment of A. a.
albinucJia best reflects its status.

Atlapetes pallidinucha

Range. — Occurs in the Eastern Andes of
Colombia, barely extending into Venezuela,
and through the Central Andes southward
to central Ecuador (Fig. 5). In Ecuador
this species seems to be almost absent on
the western slope although there are many
records from the eastern slope.

A. paUidinucha is principally a temperate
region species, although on occasions it de-
scends to the upper subtropical zone and
ascends into the paramo zone. No other
atlapetes is found in the paramo. It ranges
from about 1,700 to 3,800 m, but most com-
monly from 2,000 to 3,100 m.

Habitat. — Olivares (1969) says the spe-
cies is found in scrub and the remnants of
forest. R. Webster ( in litt. ) noted the spe-
cies sympatric with A. rtifinucha in moist
shrubbery and in forest edges in Ecuador.
It is also sympatric with A. torquatus in
similar habitats around Bogota ( Webster, in
litt.; Donahue, verb. comm.).

//a/?/^.?.— Although Olivares (1969) found
A. paUidinucha to be the most abundant
"fringillid" in Cundinamarca, and the num-
ber of records from elsewhere also suggest
it is a common species, nothing seems to
have been recorded of its habits or voice.

Morphological variation. — Females have
slightly shorter wings and tails than the
males.

' paUidinucha
papallactae

Figure 5. Distribution of A. paUidinucha.

Two races are recognized, the nominate
form from the Eastern Cordillera of Colom-
bia and adjacent Venezuela, and A. p.
papallactae, a somewhat darker race, from
the remainder of the range.

Atlapetes rufinucha superspecies

A. melanocephalus, which is endemic to
the Sierra Nevada de Santa Marta, Colom-
bia, seems to have been derived recently
from nearby A. rufinucha. The two taxa
are, therefore, treated as allospecies.

Atlapetes rufinucha

Range. — A. rufinucJia has a wide but
disjunct distribution in the Andes (Figs. 6,
7 ) . It reaches its northernmost point in the

Biology and Evolution of Atlapetes • Paynter 331

Figure 6. Northern races of A. rufinucha. A. r. sim-
plex is recorded from "Bogota" but probably comes
from farther south in the Eastern Andes.

Sierra de Perija along the Colombia-Vene-
zuela border (A. r. phelpsi). It next occurs
in three widely separated localities in cen-
tral Colombia, viz., near the northern end
of the Central Andes (A. r. eloeoprorus),
possibly in the central or southern portion
of the Eastern Cordillera although it is
known only from native "Bogota" specimens
(A. r. simplex), and in south-central Co-

melanolaemus

rufinucha

carrikeri

Figure 7. Southern races of A. rufinucha.

loinbia on the western slope of the Centi'al
Andes and on the eastern slope of the West-
ern Andes ( A. r. cauccie ) .

The species then occurs, more or less con-
tinuously, from Narifio, southern Colombia,
south on the Pacific slope and interandean
plateau through Ecuador to northwestern
Peru (A. r. spodionotiis south to central
Ecuador where it intergrades with A. r.
coniptus of farther south). On the Ama-
zonian slope of Ecuador there are records
from the north in the vicinity of Papallata,
but for the next 250 kilometers the species
is absent; it reappears in Azuay, as the race
latimicluis, and is common from here south-
ward well into Peru. It is almost certain
that the distributional gap is real and not
a collecting artifact; several areas within
the gap have been well-collected ( see Payn-
ter and Traylor, 1977:138).

In northern Peru A. r. comptus occurs on
the western slope in Piura and is replaced
farther south by A. /•. chugurensis. On the
eastern slope A. r. latinuchus reaches Ama-
zonas and A. r. J)aroni is in southern Caja-
marca and Libertad, just entering the Pa-
cific di-ainage west of the upper reaches of

332 Bulletin Miisemu of Comparative Zoology, Vol. 148, No. 7

the tributaries of tlie Maraiion west of
Cajamarca. The species is found no farther
south on the western slope l)ut recurs on
the Amazonian slope in southern Peru ( Fig.
7) in Cuzco and Puno (A. r. melanolaemus)
continuing through eastern Bolivia (nomi-
nate rufinucha) to Santa Cruz (A. r. ccir-
rikeri). \Miile the gap in distribution in
eastern Ecuador seems genuine, the gap in
eastern Peru is less certain, owing to the
paucity of collections from that region.

It is a species of subtropical and, less
often, temperate regions. It has a notably
wide altitudinal distribution, ranging from
as low as 600 m in Bolivia (nominate
rufinucha; range 600-3,350 m) to as high
as 3,700 m in southernmost Colombia ( A. r.
spodionottis; range 1,760-3,700 m). While
there seems to be a tendency for the species
to occur at generally higher elevations near
the equator, the data are too scanty to con-
firm this. It is probable that the equatorial
population will be found to average only a
few hundred meters higher than those popu-
lations at the ends of the range.

Habitat. — Mainly in thick mesophytic
scrub of moderate height, but also found in
drier or wetter situations. It seems to be
most common in moderately wet habitats,
although near Gonzanama, Loja, in south-
ern Ecuador, I once observed it in small
numbers in a rather dry subtropical area
which was also occupied by A. nationi
simomi (Paynter, 1972:303) and there are
numerous records of the species in the dry
interandean region of northern Ecuador.

Habits. — I observed the species on several
occasions in southern Ecuador and noted it
to be somewhat less shy and more arboreal
than most species of Atlapete.s. At each ob-
servation, the species was from half a meter
above the ground to as high as six meters.
In the latter instance a small flock was feed-
ing in bromeliads and other epiphytes. The
species is notably active for an atlapetes. It
moves about in flocks of up to five or six in-
dividuals, at least when not breeding. These
observations confirm those of Taczanowski
( 1884 ) made in Peru.

Little is known of the breeding season.
No breeding was noted in a long series of
specimens collected from late July through
October in southern Ecuador ( MCZ collec-
tion). Chapman (1927) reported a male
with much enlarged testes in Cajamarca,
Peru on 20 April.

I have heard A. rufinucha give two calls.
One is a single high-pitched thin zeep,
which seems feeble for a bird of its size.
This call appears to be given when a bird is
concealed and relatively inactive. Presum-
ably it is used to communicate with other
members of the flock when they are scat-
tered and out of sight. The second call is
a series of louder, lower-pitched notes re-
sembling the calls of squabbling flycatchers,
such as Myiozetetes simili.'i, although not
nearly so loud. This call has been heard
when a flock is actively moving about;
several birds may call at once. No song has
yet been noted.

Morphological variation. — Twelve races
have been described. Most of them are ex-
tremely well-differentiated from one an-
other. A. rufinucha is, moiphologically, one
of the most variable species within the
genus, even exceeding highly variable A.
torciuatus.

No clear difference in size is evident be-
tween the various subspecies although the
few measurements that are available for the
southernmost race (A. r. carrikeri) suggest
that its wing and tail may be slightly shorter
than those of other subspecies. In contrast
to size, however, some of the variations in
color are striking.

In addition to more subtle or gradual dif-
ferences in the color of the tail, venter, etc.,
there are six characters that vary markedly
within the species (Table 1). No pattern
of variation of a single character or corre-
lation of one character with another appears
to exist. For example, a sizable alar specu-
lum is present in elaeoproriis, caucae, and
latinuchus, but all three forms are widely
separated from one another and not bridged
by either chuguren.sis or simplex, the two
forms with small (or inconsistently present)

Biology and Evolution of Atlapetes • Paynter 333

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334 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

patches. Another example is the yellow
loral spot that is present in elaeoprorus,
simplex, caucae, comptus, chuguremis,
haroni, and nominate rufinucha, all of
which, with exception of chugurensis and
haroni, are well separated from one another,
either by distributional gaps or by the
presence of taxa lacking the character.
Even melanolaemus and carrikeri, which re-
semble one another more than they do any
other races are not contiguous or even proxi-
mate, but are separated by nominate
rufiimcha which is more similar to the
northern races.

Noteworthy, because it might appear to
be a clue to evolutionary relationships, is
the fact that A. r. haroni bears a very strong
resemblance to A. pallidinucha papallacta.
However, I do not think the two are related.
The latter is larger and greener, ventrally,
with the white of the nape more restricted,
and its white feathers edged with black.

Atlapetes melanocephalus

Range. — Endemic to the Sierra Nevada
de Santa Marta, Colombia (Fig. 8) where it
occurs in the upper tropical and the sub-
tropical zones at altitudes from 600 to
2,400 m.

Ualntat. — Reported by Todd and Car-
rikcr (1922:525) to occur in "almost all
kinds of conditions throughout its range — in
the forest, scrub-growth, and even in low
bushes in the open."

Habits. — In contrast to other members
of the genus, A. melanocephalus is stated
by Todd and Carriker (1922:525) to be
"not at all shy"; they also note that it
keeps near to the ground, moves about in
pairs, and lays two white eggs in a domed
nest low in a bush. No other atlapetes is
yet known to build a domed nest and for
that reason the observation is suspect.

Atlapetes tricolor superspecies

The three species comprising this super-
species are little-known but they are, mor-
phologically, quite similar, except for the

Figure 8. Distribution of A. melanocephalus.

color of the pileum. I have no hesitancy in
postulating a monophyletic origin for them.

Atlapetes flaviceps

Range. — Known from only two specimens
collected over 65 years ago in the subtropi-
cal zone (2,050-2,160 m) at Toche and Rio
Toche in a deep valley on the Nevada de
Tolima on the eastern slope of the Central
Andes of Tolima, north-central Colombia
(Fig. 9).

Hahits. — Nothing is known of the living
bird.

Atlapetes fuscoolivaceus

Range.

-Most of the few known speci-
mens were collected at least 50 years ago.
These specimens and more recent observa-
tions indicate that the species occurs only
in the subtropical zone, at altitudes from
1,500 to 2,400 m on the eastern side of the
Central Andes in the upper Magdalena
Valley, Iluila, Colombia (Fig. 9).

Hahitat. — The four localities from which
the species is known (San Agustin, La
Palma, La Candela, and Moscopan) are
(were?) in the region of dense, tall, sub-
tropical forest. I have seen one specimen
bearing on its label th.e notation "forest."

Between 3-5 April 1977, I saw the species
three times at San Agustin. I observed
one bird in very thick, 2-meter high second-

Biology and Evolution of Atlapetes • Paijnter 335

growth; the second was in a hedgerow with
dense brush and small trees up to 5 meters
in heiglit, and the third was in a row of 15-
meter trees whose tops were only slightly
above the level of a road cut into the side
of the hill. Although in a region of high
rainfall, the sites where the birds were
noted were not particularly lush. There re-
main patches of fairly high forest around
the archaeological sites of San Agustin, but
no atlapetes were seen within the forest.

Habits. — Only single birds were seen at
San Agustin. No calls were given, but
once the beat of the wings, a characteristic
noted in several other forms of Atlapetes,
was heard as the bird flew off.

Two of the three birds seen were surpris-
ingly high in the vegetation. One was two
meters above the ground and easily seen
against the horizon, but it quickly dropped
down when approached. The other was in
the crown of a 15-meter tree, where it was
leisurely feeding on dark purple or black
fruit about 5 mm in diameter in company
with various tanagers, warblers, and a
Swainson's thrush (Cafharus iisttdafiis) .
The bird was watched for five minutes until
it dropped down into shrubs and disap-
peared. The rich yellow underparts, and
even the streaked throats, were conspicuous
on these two birds whose behavior was in
such contrast to the usual furtive demeanor
of other atlapetes.

Chapman ( 1914 ) reported the type spec-
imen, a male from San Agustin, had much
enlarged gonads. The specimen probably
was collected in April or May (see Chap-
man, 1917:45).

Atlapetes tricolor

Range. — The species is known in Colom-
bia on the eastern slope of the Western
Andes from a single specimen ( San An-
tonio, Valle) and on the western slope of
the same range from several specimens
taken from Caldas south to Nariiio, with the
majority of reports from the latter depart-
ment. Although San Antonio is the type
locality for the species (specimen taken by

A. flaviceps
A. f uscooli vaceus
A. \. crassus
At. tricolor

▲

Figure 9. Distribution of A. tricolor superspecies.

M. G. Palmer, 11 Dec. 1907, according to
Bangs, 1908:61), the absence of any other
record from the eastern slope of the \\'est-
ern Andes makes the record suspect. Fur-
thermore, in a year spent at San Antonio in
1958-59, Miller (1963) failed to find this
species. There is a cluster of records from

336 Bulletin Museum of Comparative Zooloiitj. Vol. 148, No. 7

adjacent northwestern Ecuador and a single
specimen, far to the soutli, from La Chonta,
El Oro, in southwestern Ecuador, again on
the western slope. The species is absent in
northern Peru but reappears in central Peru
on the eastern slope in San Martin and
ranges south to Cuzco (Fig. 9).

The species seems to have a particularly
wide altitudinal range in the northern por-
tion of its distribution, occurring from some-
what below 300 m up to 2,000 m and from
the moist upper tropical zone to the sub-
tropics. In Peru, however, it seems not to
occur much below 1,700 m and ranges up to
at least 2,400 m which means that in this
region it is a species of the subtropical and
lower temperate zones.

Habitat. — Taczanowski (1884) quoted
Jelski's observation that the bird frequented
dense thickets. Terborgh (in litt. ) has seen
it in Peru in elfin forest which was barely
shoulder high and in undergrowth border-
ing small clearings. R. Webster (in litt.)
saw the species around Tandapi [== Manuel
Cornejo Astorga; 00 "25'S/78^48'W], Ecua-
dor in dense thickets within the forest and
on its edges.

Habits. — This atlapetes seems to be rare.
It is little-known; Jelski observed it in Peru
nearly a century ago (Taczanowski, 1884)
and reported it to be a shy bird that moves
about in small bands, but becomes more
bold and more visible when courtship and
song begin in February.

Richard Webster (in litt.) saw the species
in June and September 1975 at three places
on the slopes between Tandapi and above
Santo Domingo de los Colorados, Ecuador.
On 19 June at Tandapi he saw three pairs in
an hour and a half; one pair was feeding a
fledgling. Webster describes the species as
being noisy, with a call of "a squeak fol-
lowed by two chirps," many single squeaks,
and a song of "siceet [slight pause], churr."
In early September, he noted the species in
a mixed flock of Synallaxis- azara, Myiody-
nastes miniatus and M. chrysocephalus,
Pipraeidea melanonota, and Tan^iara ni<ji^ro-
viricUs: Jelski (Taczanowski, 1884) re-

ported that flocks of A. tricolor are
sometimes accompanied by one or two in-
dividuals of A. schistacetis.

Morphological variation. — The northern
and southern populations, which are widely
separated, have been recognized as racially
distinct. The northern race (crassus) has a
noticeably thicker and longer bill and is
generally darker and more richly colored.

de Schauensee ( 1951 ) thought that there
might be sufficient variation in the northern
taxon to warrant naming additional races.
This led Miller (1960) to believe crassus
might even be a distinct species, but de
Schauensee had based his speculations on
the mistaken belief that the type of crassus
has a brown crown, when it is instead
merely a rich, tawny gold, not differing ap-
preciably from other specimens from Co-
lombia and Ecuador.

Atlapetes albofrenatus

Range. — Occurs in the Eastern Andes of
Colombia from about Bogota northward to
Santander and in the Andes of Venezuela
in Tik-hira and Merida (Fig. 10). Its alti-
tudinal range is from about 1,000 to 2,500
m, but it seems mainly to occur above 1,600
m, and thus most often inhabits the sub-
tropical zone and less frequently the upper
tropical zone.

Habitat. — I saw the species on two occa-
sions, 1-2 April 1975, in thick subtropical
cloud forest, as well as in more open scrub,
a short distance north of Bucamaranga, Co-
lombia, at altitudes of 2,000-2,400 m. The
birds moved into somewhat exposed situa-
tions for brief periods but preferred dense
tangles such as the masses of ferns and
bamboo that cascade over road cuts. Phelps
and Phelps, Jr. (1963) imply that in Vene-
zuela it is a species of forests. Webster (in
litt. ) informs me that he saw this atlapetes
at altitudes ranging from 2,100 to 2,300 m,
in dry, thorny, low scrub, on tlie mountains
above Villa de Leiva, Boyaca, Colombia.
The species' tolerance of a wide range of
vegetation is confirmed by Olivares who
wrote ( in litt. ) that the bird "occurs in low

Biology and Evolution of Atlapetes • Paynter 337

meridae

albof rena t

•

y^,^

r-<^

/.> ~\^

/ 0

\ /

)

V \ ^

)

•

{

V*.

Figure 10. Distribution of A. albofrenatus.

Figure 11. Distribution of A. leucopis.

forest which ranges from dry to moist, as
well as in very humid subtropical forest."

Habits. — North of Bucamaranga the spe-
cies was seen singly, paired, and in what
appeared to be small family groups. They
were mainly close to the ground but at times
10 m high in trees. In general their move-
ments were, conspicuously, more rapid than
those of A. schistaceiis, which was also
noted in the same area. The onlv call heard
was a thin "zeep," characteristic of the
genus. Webster (in litt.) also noted the spe-
cies' propensity to range high in trees in the
scrubby oaks bordering streams on the dry
slopes above Villa de Leiva. Olivares (in
litt.) has found the species in the lower
branches of trees in the interior of the
forest and says that an examination of sev-
eral stomachs revealed mainly insects in
some, while others contained more seeds.
I observed a bird catch a small white moth.

Morphological variation. — Two well-dif-
ferentiated races are recognized. The race
meridae, of the Merida Andes, which is
separated from nominate allyofrenatus of
the Eastern Andes of northern Colombia by

the valley of the Rio Torbes, is markedly
different in having reduced malar stripes,
a forehead that is concolor with the crown
instead of being black, and underparts that
are entirely yellow rather than white on the
throat and upper breast. No difference in
size is apparent.

Atlapetes leucopis

Range. — Known with certainty from only
three localities, apparently all in the sub-
tropical zone (Fig. 11). The first is at La
Plata (2,350 m) which is in a deep valley
on the eastern slope at the southern end of
the Central Andes and at the head of the
Magdalena Valley in Huila, Colombia. The
second is on Cerro Pax ( alt. ? ) , which is on
the eastern slope in Nariiio, also in southern
Colombia. The third is at Palmas (ca.
2,500 m), Azuay, on the eastern slope of
southeastern Ecuador. The species was de-
scribed from "Yauayaca" [= ? Yanayacu,
fide Paynter and Traylor, 1977], a locality
somewhere on the east slope of Ecuador.

Habitat. — Unknown.

Habits. — Only eight specimens are be-

338 Bulletin Muscinn of Comparative Zoology, Vol. 148, No. 7

Figure 12. Distribution of A. pileatus.

lieved to exist (de Schauensee, 1951). It is
among the least known of all atlapetes.
There are no accounts of the living bird.

Morphological variation. — de Schauensee
(1951) noted that a single bird from La
Plata differed in color from four specimens
from Cerro Pax, but there is no way to ap-
praise the significance of this single obser-
vation.

Atlapetes pileatus

Range. — Distributed on the Mexican Pla-
teau from Chihuahua and Tamaulipas south
to Oaxaca (Fig. 12). Occurs from 900 to
3,500 m, but most commonly from 1,500 to
2,800 m, in the temperate zone.

Habitat. — Moderately moist undergrowth,
usually at edges of pine, oak, or pine-oak
forest.

Habits. — Little is known of the habits of
this secretive bird, which does not seem to
be particularly common anywhere.

It has been described as remaining within

a few feet of the ground, moving rapidly,
being "nervous," and jerking its tail (Ed-
wards, 1972). Cody and Brown (1970) re-
port that 90 percent of the bird's foraging
is done on the outer branches of small trees
and bushes and that, at least during the
breeding season in Oaxaca (late March-
early April), the species seems to be in-
sectivorous. There is no further information
on feeding habits.

The breeding season is prolonged, extend-
ing from late April in Oaxaca (Cody and
Brown, 1970) to the end of August in San
Luis Potosi ( Miller et al., 1957) .

The voice, which is "thin" and delicate,
has been described by Edwards ( 1972:257)
as "a high-pitched wees, a metallic chip, a
chip-chip-chip chatter; and several song
variations of a series of notes, such as chip-
ee-wee-wee-wee or chip-ee-r-r — r-r-r, as-
cending in pitch to the first icee or r-r note."

Morphological variation. — The northern
population (A. p. dilutm) is slightly smaller
and generally paler than the nominate form.

Biology and Evolution of Atlapetes • Paynter 339

Figure 13. Distribution of A. semirufus and A. personatus.

These differences are easily distinguished
but of no great magnitude. It is noteworthy
that this is one of the few atlapetes exhibit-
ing intraspecific variation in size.

Atlapetes fulviceps superspecies

This superspecies consists of three al-
lospecies, viz. semirufus, personatus, and
fulviceps. Although widely separated geo-
graphically (Figs. 13, 14), the three are
morphologically very similar, \\\\\\ chestnut
heads, yellow underparts, and green backs.
Chapman, as long ago as 1931, postulated
that personatus and fulviceps might be con-
specific; this may be the eventual dispo-
sition of all three taxa, l)ut I should like to

know more of the living birds before taking
this step.

Atlapetes semirufus

Range. — Occurs from the coastal Cordil-
lera of northern Venezuela southward
through the mountains to Cundinamarca in
the Eastern Andes of Colombia (Fig. 13). At
the northeastern extremity of its range the
species ( A. s. denisei ) descends to as low as
600 m in the upper tropical zone and as-
cends as high as 2,100 m in the subtropical
zone. Elsewhere the species does not occur
below 1,200 m and is in the subtropical
zone, exclusively, except at its southern-
most limits where in the vicinitv of Choachi,

340 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

Figure 14. Distribution of A. fulviceps.

Cundinamarca it ranges Ijetween 3,000 and
3,500 m and is in the tenipeiate/subparamo
zone (Olivares, 1969).

Habitat.— Schiiier and Phelps ( 1954 ) re-
ported that A. s. denisei has a wide ecologi-
cal range, occurring in moderately moist to
somewhat xerophytic second-growth and in
the weeds of low forest. Nothing seems to
have been published concerning the habi-
tat requirements of the other races. Because
they appear to be confined to the subtropi-
cal zone (or temperate zone in the case of
nominate semirufus), they probably have
more restricted habitat preferences.

Habits. — Again, all we know about this
species is contained in the brief account by
Schiifer and Phelps ( 1954 ) . They found
that at Rancho Grande, Aragua, Venezuela
the bird is of limited abundance, sedentary,
and occurs in pairs or in groups of up to

eight individuals. Breeding is from May to

July.

Schiifer and Phelps' observations suggest
that this species might benefit by the crea-
tion of second-growth through human activ-
ity; one might even imagine the bird be-
coming more abundant than it was in the
past. However, Olivares ( 1969 ) observes
that in Cundinamarca, it was common early
in this century, and he implies that there-
after it was exceedingly rare until a number
of birds were discovered at Une in 1967.
Whether this is an indication of a recovery
of the species as a whole or merely the for-
tuitous discovery of a restricted population
is imknown.

Morplwlugical variation. — Morphological
variation is not great, although six races
have been recognized. Geographical varia-
tion in the shade of green on the back and of
the brownish rufous and yellow on the
venter are the obvious differences. There
may be variations in size, but if so, they are
of no great magnitude as they are not de-
tectable in the limited material available
for study.

Atlapetes personatus

Ran<s,e. — Found in scattered and isolated
populations in the moist subtropical zone,
between 1,0U0 and 2,500 m, on the table
mountains ( "tepuis" ) of southern Venezuela
and adjacent Brazil (Fig. 13).

Hal)itat. — From the descriptions of Chap-
man (1931) it appears that the species oc-
curs in the brushy wet forests on the sum-
mits of the tepuis.

Hahits. — Nothing recorded.

Morphological variation. — The species
has been divided into six subspecies, most
of which are well-differentiated by color,
pattern, or both. There are two basic color
patterns, viz., that in which the bird's entire
head, chin, throat, and chest are chestnut
and that in which the chestnut does not
extend to the throat and chest. Nominate
personatus and A. p. collaris occur in south-
western Bolivar in relatively close proxim-
ity. They are of the latter type and the re-

Biology and Evolution of Atlapetes • Paynter 341

maining four races, whicli lie to the west,
have the former pattern. The two color pat-
terns correspond to the two zoogeographical
subdivisions of the "pantepui" region which
lie on either side of the Rio Cavn'a ( Mayr
and Phelps, 1967). Variation within these
two groupings is in the extent of the chest-
nut color or in the general intensity or shade
of the colors. No significant differences in
size have been noted.

Atlapetes fulviceps

Range. — This little-known s]3ecies has
been recorded from the eastern slopes of
Bolivia (La Paz to Chuquisaca) and in
nt)rthwestern Argentina, apparently from
about 400 m to as high as 2,700 m ( Fig. 14) .
Hellmayr ( 1938) states that this is a form of
the subtropical zone, bvit if the altitudinal
range is as wide as the collecting records
seem to indicate, the species must reach
the temperate, or at least subtemperate,
zone. Apparently this is a very rare bird
in Argentina.

Ha])itat and habits. — Nothing recorded.

Morphological variation. — No geographi-
cal variation in morphology has been de-
tected.

Atlapetes citrlnellus

Range. — The species, which is the south-
ernmost representative of the genus, is re-
stricted to the subtropical slopes of the
Andes in Salta, Jujuy, and Tucuman, north-
western Argentina, from about 1,000 to
3,100 m (Fig. 15). I can find no substantia-
tion for Olrog's ( 1963 ) inclusion of Cata-
marca within the range. There is one sus-
pect record from northern Paraguay ( Ber-
toni, 1924).

Ha/;jfaf.— Dinelli (1918) reported that
the species occurs in dense ferns in shady,
humid ravines; Wetmore ( 1926) found it in
thick weeds at the edge of a grove.

Hfl/;/Y.s.— Dinelli (1918) noted this bird
on the ground in small groups and reported
it is not timid and is readily lured into sight
when its call is imitated (described as a
faint "t.sip" by ^^'etmol•e, 1926). This is a

Figure 15. Distribution of A. citrinellus.

remarkably different type of behavior for
an atlapetes. Dinelli (1918) describes the
nest as being about 1^2 m above the ground
in a shrub. The egg is rosy-pink or white
with dark brown or maroon spots and
blotches, principally at the large end (Di-
nelli, 1918; Smyth, 1928). Smyth (1928)
implies that a clutch consists of three eggs,
and Percy ra ( 1951 ) also mentions a nest
with three eggs; the latter was collected on
21 November, which is the only breeding
date available for this little-known bird.
Unfortunately, there appears to be no de-
scription of the nest itself.

Morphological variation. — There is no in-
dication of either sexual or geographical dif-
ferences in moq3hology.

Atlapetes brunneinucha

Range. — A. brunneinucha ranges from
northwestern Mexico to southeastern Peru
(Figs. 16, 17) and thus has the most exten-
sive distribution of any atlapetes. It is a
species of humid regions, generally occur-
ring in the subtropical zone although at
times it ranges well down into the tropical
zone or up to the lower temperate zone,
but only where the vegetation is lush and
dense, such as in cloud forest.

342 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

brunneinucha

•

apertus

suttoni

macrou ru s

alleni

elsae

Figure 16. Northern races of A. brunneinucha.

At the northern end of its range, the spe-
cies occurs as low as 200 m ( nominate
brunneinucha of eastern Mexico) and 350
m (A. h. apertus of the Sierra de Tuxtla,
southern Veracruz) but on the other hand
in the same area it also reaches an extreme
altitude of 3,500 m (A. /;. suttoni of Guer-
rero and Oaxaca ) .

This is doubtless the maximum altitudinal
range for any species of atlapetes within a
reasonably circumscribed area. In Central
America, the species ranges from 550 to
3,400 m and in South America from 900 to
3,400 m. Throughout, however, it seems

most abundant between roughly 1,000 and
2,000 m.

Because of the abundance of moist sub-
tropical forest, the species is widely dis-
tributed with few major discontinuities.
As with birds of similar requirements, the
species is absent in most of Panama, but it
occurs throughout the Andes south to
northern Peru where, owing to the absence
of moist forests, it drops out on the western
slope but continues southward on the east-
ern side.

In the coastal range of northern Vene-
zuela there is an interesting transposition of

Biology and Evolution of Atlapetes • Paynter 343

f ron ta lis
allinornatus
inorna t us

Figure 17. Southern races of A. brunneinucha.

344 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

relative altitudes between A. b. frontalis and
A. torquatiis phaeopleurus. Throughout
their joint ranges, A. torquatus occupies the
higher altitude and A. brunneinucha the
lower, but at Rancho Grande, Aragua, A.
torquatus ranges from 700 to 900 m while A.
brunneinucha occurs from 900 to 2,400 m
but its optimum range is between 1,600 and
2,000 m (Schafer and Phelps, 1954).

Habitat. — The bird is an inhabitant of
the interior of humid forest or of the thick
undergrowth at the edges of the forest, but
is still under taller trees. This propensity
for the cover of forests has been remarked
upon by Miller (1963) and Slud (1964).
The latter also pointed out tliat in Costa
Rica A. gutturalis [= albinucha] and A.
brunneinucha have ranges in common but
A. albinucha is a nonforest species. This
dichotomy in habitats seems to hold
throughout the ranges but no broadening
of the habitat of A. brunneinucha seems to
occur south of Colombia where A. albinucha
is absent.

Habits. — The ecology of A. brunneinucha
is fairly well known considering the diffi-
culty in obtaining this information for other
members of the genus. This is perhaps
because, although a quiet species, it does
not seem to be so shy as some other atla-
petes.

As was noted long ago by Taczanowski
(1SS4), and later by Carriker (1910), A.
brunneinucha is usually found in pairs or
in family groups. Schafer and Phelps (1954)
reported a maximum flock size of six.

A. brunneinucha is a bird of the dark
forest floor, although Miller ( 1963 ) once
found it 20 feet up in second-growth where
it was feeding on seeds in company with
tanagers. This, however, is probably an in-
frequent practice. Miller also comments on
the bird's habit of kicking dry leaves in
search of food, although Taczanowski (1884)
reported that the species tossed the leaves
with its beak. While in Venezuela and
Peru, Fitzpatrick ( in litt. ) has seen the bird
stir the litter with its feet but has never
seen it use its bill for this purpose.

Miller ( 1963) believed that, in Colombia,
breeding probably occurs throughout the
year. Farther north, in Mexico and Central
America, breeding seems to begin in March
or as early as late February and extends to
April or Mav or even June ( Miller et al.,
1957; Blake, 1958; Carriker, 1910). Breeding
in Venezuela has been recorded from April
to June ( Schafer and Phelps, 1954 ) ; in Peru
the only breeding record known to me is
that of a nest found in January ( Taczanow-
ski, 1884).

The nest, which is placed in shrubs or
ferns close to the ground, is made of coarse
leaves and lined with finer material (Tac-
zanowski, 1884; Carriker, 1910). The clutch
seems to consist of but one or two greenish
or bluish white eggs (Sclater and Salvin,
1879; Carriker, 1910).

A. brunneinucha appears to be omnivo-
rous (Olivares, 1969), although Taczanow-
ski ( 1884) examined one specimen that had
eaten only invertebrates.

As with all forms of Atlapetes, this species
has an inconspicuous voice. Taczanowski
(1884) described its call note as a series of
rapid tsit-tsit notes and its song as reminis-
cent of the "voice" produced by a rubber
doll; presumably this refers to "a series of
high-pitched notes" (Peterson, 1973). Mil-
ler ( 1963 ) , in spite of long-term observa-
tions in Colombia, never heard a song but
described the infrequently produced alarm
calls as chattering tsip's and noted a mewing
note suggesting that of Pipilo chlorurus.

Morphological variation. — Chapman
( 1923a ) in his review of the genus Buar-
remon [= Atlapetes] pointed out that A.
brunneinucha probably has a more exten-
sive range than any other species of the
subtropics and believed it exhibited no geo-
graphical morphological variation in spite
of its wide distribution. Chapman did,
however, recognize the taxon inornatus as
being close to brunneinucha but, as was
customary at that time, treated it as a full
species rather than as a race of A. brun-
neinucha. Parkes (1954) in his review of
the species detected more variation and rec-

Biology and Evolutiox of Atlapetes • Paynter 345

ognized nine races, five of which he de-
scribed. Subseqiientlv two more races were
named (Phillips, 1966; Rowley, 1968)
bringing the total to 11 subspecies. There
is no doubt that there is geographical vari-
ation within the species, but most of this
variation is extremely subtle and probably
not worth nomenclatural recognition.

There is some geographical variation in
size (see Parkes, 1954, p. 134 for details).
A. b. frontalis, the most widespread race of
South America, has a slightly longer bill
than all other populations, possibly except-
ing the races inornatus and aUinornatus
which are restricted to west-central Ecuador
and central Venezuela, respectively. A. b.
suttoni of Guerrero and Oaxaca and particu-
larly A. b. macrotirus of Chiapas and Guate-
mala have generally longer tails than the
remaining populations. No geographical
trend in these variations is evident.

Variations in color and patterns are also
not pronounced. The extent of yellow bor-
dering the chestnut crown varies somewhat,
as does the amount (jf black on the forehead
and what portion of the venter is occupied
by either gray or white. Presumably these
variations are of no biological significance.
There is one characteristic, however, that
does have a pattern, although its signifi-
cance is also not apparent. This is the ab-
sence of a black breast band in three small,
isolated or relatively isolated populations.
One of the characteristic and striking mark-
ings of most taxa of A. brunneinucha is a
black band that separates the white throat
from the gray, or gray and white, chest and
abdomen. In A. b. apertiis, which is isolated
in the Sierra de Tuxtla of Veracruz, Mexico,
in A. /;. aUinornatus of the Sierra de San
Luis and Sierra de Aroa in northwestern
Venezuela, and in A. b. inornatus which
occurs in a pocket on the western slopes of
the Andes of central Ecuador, the black
pectoral band is absent or rudimentary. The
significance of this is unknown. The isola-
tion of the three taxa leads one to suspect
that the breast band is a species recognition
character which might be unnecessary in

small or isolated populations. A. h. apertus
lacks congeners nearby but both A. h. aUin-
ornatus and A. 1). inornatus do have congen-
ers in the vicinity. Some relationship be-
tween the lack of a breast band and the
presence of closely allied A. torquatus also
suggests itself, but A. torquatus, while near
A. /;. inornatus, is absent from the range of
A. b. apertus and A. /;. aUinornatus. Further-
more, in Mexico and most of Central Amer-
ica, where A. torquatus is absent, A. ]}run-
neimicha has a breast band. Especially
baffling is the fact that the width of the
band does not vary appreciably between
the populations that do possess it, which is
contrary to what one might expect if this is
especially important in inter- or intraspecific
recognition in large populations. Parkes
( 1954 ) has pointed out that there is no way
to resolve whether the banded or unhanded
condition is ancestral to the other.

Atlapetes torquatus superspecies

There are 15 allopatric taxa that are very
similar morphologically, that certainly are
closely related, and that doubtless are best
treated as members of a superspecies, viz.,
A. torquatus. The torquatus superspecies
seems to be comprised of the allospecies
virenticeps, atricapiUus, and torquatus, but
to which allospecies certain of the 15 taxa
belong is a \'exing problem, a circumstance
not met with elsewhere in the genus. The
root of the problem is the rather major mor-
phological variation between a number of
the 15 taxa, further complicated by the fact
that variation occurs within several char-
acters, and these \'ariations are seldom con-
cordant.

Except for the absence of a chestnut
pileum, most forms of A. torquatus super-
species are notably similar to A. brun-
neinucha, a species which has about as ex-
tensixe a range but which exhibits little
geographic variation.

Within the A. torquatus superspecies
there is some minor geographical \ariation
in overall size and in the relative lengths of
the wing and tail, and somewhat greater

346 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

Figure 18. Distribution of A. virenticeps, of tfie tVlexi-
can plateau.

variation in the color of the venter, but
larger differences occur in the pattern and
color of the head, in the presence or absence
of a pectoral band, and in the color and
extent of the superciliary stripe. Two forms,
tacarcunae and atricapilhis, have markedly
more robust bills.

There are three basic head types, viz. all
black {tacarcunae and atricapilhis), black
with a medial stripe and superciliaries that
are yellowish green ( virenticeps ) , and black
with a medial stripe and superciliaries that
are either gray or white (the remaining
12 taxa). The geographical distribution of
these three patterns is relatively simple
(Fig. 20); virenticeps is in Mexico, tacar-
cunae and atricapilhis are in eastern Pan-
ama and north-central Colombia, respec-
tively, and the other forms occur, some in
isolation and others contiguously, from
Costa Rica to Argentina.

The distribution of taxa with or without
breast bands is more complex (Fig. 20).
The forms virenticeps, costaricensis, tacar-
cunae, atricapilhis, assimilis, nigrifrons, and
borelli lack the band; the remaining eight

taxa have it. In geographical terms this
means that forms without breast bands oc-
cur in the northern part of the range, except
for the Sierra Nevada de Santa Marta and
the V^enezuelan Andes, and in the extreme
south, while forms with pectoral bands oc-
cur in the southern range, with the excep-
tion of southern Bolivia and northwestern
Argentina which are occupied by borelli, a
ta.xon without a band.

The color of the superciliaries ( ignoring
minor departures from these generalizations,
such as white lores with gray stripes) is
either ( a ) black ( = absent ) , which is the
case in black-headed tacarcunae and atri-
capiUus, or (b) yellowish-green, as in viren-
ticeps, or ( c ) gray, as in costaricensis, basili-
cus, perijamis, larensis, assimilis, nigrifrons
and poliopJirys, or (d) white, which is the
condition in the five taxa remaining. The
geograpliical pattern (Fig. 20) is even more
complex than that which exists for pectoral
bands. Agreeing with the pattern of distri-
bution found for head color are virenticeps
and also tacarcunae and atricapilhis. White
eye stripes occur in the outliers at both ends
of the Andes, viz. phaeopleurus and phijgas
in north and northeastern Venezuela and
torquatus, fimbriatus, and borelli in Bolivia
and northwestern Argentina. The remain-
ing taxa have gray stripes. From this distri-
bution one might suspect the existence of
some sort of relationsliip between outlying
populations and white superciliaries, but
this is dispelled upon recalling that basilicus
of the Sierra Nevada de Santa Marta and
costaricensis of Costa Rica and westernmost
Panama are both completely isolated but
have gray eye stripes. There exists no cor-
relation between breast band and the color
of the superciliaries; all five forms with
white superciliaries have breast bands but
of the seven with gray eye stripes, four have
pectoral bands and three do not.

Chapman (1923b) was the first to at-
tempt to determine the interrelationships of
this perplexing complex of birds. However,
this work was done when "mutations" had
attracted the fancy of biologists and when

Biology and Evolution of Atlapetes • Faijnter 347

many phenomena were attributed to "mu-
tations." We now know, however, that these
phenomena have been brought about by a
variety of less radical causes. Chapman,
dealing with 13 taxa ( perijanus and larensis
had not been named at that time), treated
the thick-billed, black-headed forms atrica-
piUufi and tacarcunae as conspecific, placed
into one species the three taxa with no
pectoral bands and gray superciliaries {as-
shnUis, nigrifrons, and costaricensis) , and
treated each of the remaining forms as a
full species. The later action was doubtless
because of the prevailing belief that even
obviously related taxa were to be considered
I distinct species unless morphological inter-
) gradation could be demonstrated.

Hellmayr (1938) maintained the two
black-headed forms as one species (A.
atricapillus) but lumped all other taxa into
a single species, A. torquatus. He even in-
cluded virenticeps of Mexico and pointed
out that it was a distinctive form that
closely resembled the immature of the more
southern taxa.

The next to deal with the problem were
de Schauensee and Eisenmann (de Schau-
ensee, 1966 ) who proposed a quite different
treatment. This made Mexican virenticeps a
full species and costaricensis a race of A.
atricapillus on the premise that tacarcunae
of eastern Panama is moiphologically inter-
mediate between costaricensis and nominate
atricapiUus. All other forms were placed in
A. torquatus. On geograpliical grounds this
treatment is appealing, because it avoids
the problem of explaining the distant isola-
tion of costaricensis from other members of
its species. Nevertheless, I cannot appreci-
ate how tacarcunae can be considered to be
morphologically intermediate between cos-
taricensis and atricapiUus. On the contrary,
I find tacarcunae difficult to distinguish
from atricapiUus and to be well-differenti-
ated from costaricensis.

The last attempt to resolve the problem
(Paynter, 1970) resulted in the lumping of
all taxa into a single species (torquatus).
The reasoning behind this was based par-

tially on an uncritical acceptance of de
Schauensee and Eisenmann's claim that
tacarcunae was intermediate between co.s-
taricensis and atricapiUus and partly on the
belief that atricapiUus and assiniiUs were
geograpliically well separated. As explained
above, costaricensis does not appear partic-
ularly close to tacarcunae. Also, since my
earlier analysis, atricapiUus and assimiUs
have been found (Olivares, 1969) within
about 25 kilometers of one another, which
is so close that it now seems a definite possi-
bility that they may be parapatric.

I think that the best treatment is to recog-
nize three allospecies within the superspe-
cies A. torquatus. A. virenticeps is the first
species. It appears to be a relict form which
in its isolation has reverted to or (less Hkely)
has retained some juvenile characteristics.

The second allospecies is A. atricapiUus,
with tacarcunae as a subspecies. Its com-
pletely dark head and thick bill, its rela-
tively restricted range, its lower altitude
(see below), and its apparent parapatry
with assimiUs suggest that this is an offshoot
of the third allospecies, A. torquatus. A.
atricapiUus seems to have differentiated
from its stem stock but has not progressed
far enough to allow it to be ecologically
compatible with A. torquatus or possibly to
be reproductively isolated from it.

Into the third allospecies, A. torquatus,
I place all the remaining taxa. The presence
or absence of a breast band and the color
of the superciliaries are probably characters
that readily respond to isolation. They may
have no adaptive significance, or they may
be important in interspecific recognition.
I am inclined to believe they are of little
significance since, for example, one can
see in the geographical continuum of tor-
quatus, finibriatus, and horeUi successive
stages in the loss of the pectoral band.

Atlapetes virenticeps

Range. — Occurs in the temperate zone of
the southwestern portion of the Mexican
Plateau and Pacific slope from Sinaloa
south to Michoacan ( Fig. 18 ) . In the north

348 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

A. a. tacarcunae

atricapillus

t. ba silic us

pe r i ja n us

la ren sis

phaeopleurus

phygas

€

assimili s

n i g r i f ro n s

Figure 19. Distribution of A. atricapillus and the centrally located races of A. torquatus. Birds from tfie West-
ern Andes have not been examined but are presumed to be referable to nominate atricapillus.

Biology and Evolution of Atlapetes • Payntcr 349

A. t. phygas

A. virenticeps

costaricensis

r y^

A. a. atricapillus

A. t.\ perijanus

A. t. larensis

A. t. assimilij

A. t. nigrifrons

A. t. poliophrys

A. t. fimbriatus ' j>

A. t. borelli Y

Figure 20. Color and pattern in the head and breast of the A. torquatus superspecies.

350 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

it ranges from about 1,100 to 1,900 in while
farther south it occurs from about 2,000 to
3,600 m.

Habitat. — Thick undergrowth of oak or
coniferous forests and at their edges (Ed-
wards, 1972).

Habits. — Ver\- Httle is known of the spe-
cies. Schaldach ( 1963 ) reported behavior
similar to that of towhees {Pipilo) in that
the bird feeds in litter, turning ox'er leaves
in its search for food. Unfortunately, he
was unable to determine whether the bill or
feet were used for this activity.

Nesting and laying have been j-joticed in
June and Julv (Miller et al., 1957; Schal-
dach, 1963). Schaldach (1963) beheved the
species to be doubled-brooded.

Morphological variation. — Some slight
variation in color and size, perhaps not
concordant with geography, seems to exist,
but the pattern described by Moore ( 1938 )
and leading to his description of a northern
race (verecundus) cannot be confirmed
( Hardy and Webber, 1975) .

Atlapetes atricapillus

Range. — The species has been found on
a few of the higher mountains of eastern
Panama (east of the Canal Zone) and in
Colombia in the middle Magdalena Valley
on the eastern slope of the Central Andes
and western slope of the Eastern Andes,
once at the northern end of the Central
Andes (above Puerto Valdivia, Antioquia)
and, recently ( Hilty, 1977) on the Pacific
slope of the Western Andes between Cali
and Buenaventura (Fig. 19). In all there
are records from only about a dozen spe-
cific localities, all of which are at an altitude
from about 700 to 1,500 m. Althoucfh vari-
ous authors (e.g.. Chapman, 1923b; Hell-
mayr, 1938) have said this is a species of
the subtropical zone, its altitudinal range
would seem to indicate it is a form of the
upper tropical zone, to which the species
was also ascribed by de Schauensee ( 1951 ) .

Habitat. — Dense undergrowth in wet
forest and edges (Ridgely, 1976).

Ha])its. — A breeding pair was collected at

La Vega, Cundinamarca (Olivares, 1969)
but the date was not recorded. Nothing
else has been published.

Morphological variation. — The birds of
Panama were separated by Chapman
( 1923b ) from the Colombian population on
the basis of slight differences in color pat-
terns and supposedly a longer, thicker bill.
The only character I am able to recognize
is a tendency for the western birds (tacar-
cunae) to have a barely distinguishable
gray postocular line, in contrast to the
solid black head of the nominate race.

The records of Hilty (1977) from the
Western Andes have been tentatively as-
signed to the nominate form, although I
have not examined the birds.

Atlapetes torquatus

Range. — The range of allospecies A. tor-
cfuatus is the most extensive of the three
taxa within the superspecies. A. t. costari-
censis, an isolate (Fig. 21), is found in
southwestern Costa Rica and presumably
in adjacent Chiriqui, Panama ( Ridgely,
1976), but I can find no specific record for
the latter. It ranges from about 1,100 m
down to 300 m in the subtropical zone and
upper reaches of the tropical zone. The
species next appears in the Sierra Nevada
de Santa Marta, Colombia, again as an iso-
late (A. t. basilicus), where it occurs from
the upper tropical zone to the temperate
zone (600 to 2,800 m) (Fig. 19).

A. t. a.ssimilis has a wide but curiously dis-
junct range (Fig. 19). In Colombia it oc-
curs on both slopes of the Eastern Andes,
but only in the vicinity of Bogota. (The
species seems to be absent south of here,
but this may be because of the lack of
observations and north of Bogota it is ab-
sent for about 300 km, until it recurs, as
other races near the Venezuelan border.)
A. t. assimilis is found on both slopes of the
mid-portion of the Central Andes from near
Medellin to about the Quindio Pass and
then reappears on the west slopes in the
vicinity of Popayan. The race also occurs
near Popayan on the eastern slopes of the

Biology and Evolution of Atlapetes • Paijnter 351

Western Andes. From the Colombia-Ecua-
dor border south to south-central Ecuador,
A. t. a.ss-imilis is found on both sides of the
Andes but in southwestern Ecuador and
northwestern Peru the race is replaced by
another (nigrifrons). In southeastern Ecua-
dor the species seems to be totally absent,
only to appear again (as assimilis) on the
eastern slope of northern Peru. Some of
the patchiness of the distribution is doubt-
less because of spotty collecting, but the
broad pattern is probably as outlined. A. t.
cissimilis has an altitudinal range from about
1,500 to 3,600 m and is typically found in
the temperate zone.

The outlying Andes of northeastern Co-
lombia and of Venezuela are occupied,
from south to north, by four races {peri-
janus, larensis, phaeopleurus, and phijgas).
They occur at altitudes from about 700 to
1,800 m in the subtropical zone and, ap-
parently on occasion (at least in phaeo-
pleurus), in the upper tropical zone (Scha-
fer and Phelps, 1954; also see discussion
under A. hrunneinuduu p. 344).

On the slopes of southwestern Ecuador
and northwestern Peru there occurs A. t.
nigrifrons with an altitudinal range from
600 to 2,700 m, which is considerably wider
than that of A. t. assimilis. It is found from
the upper tropical zone up to the beginning
of the temperate zone.

The species seems to be absent from
northern ( except the extreme north ) to cen-
tral Peru. It reappears on the eastern slopes
of central Peru and ranges from here
through eastern Bolivia to northwestern
Argentina in a series of four subspecies
{poliophnjs, torquatus, fimhriatus, and
boreUi ) . These generally occur in the tem-
perate and subtropical zones in the north
but are restricted to the subtropical
zone in the south (Fig. 22). There is, of
course, a corresponding drop in altitudinal
ranges from north to south {poliophnjs, ca.
1,800-3,650 m; torquatus, ca. 2,000-3,100 m;
fimhriatus, ca. 700-3,050 m; borelli, ca. 400-
1,200 m).

From this survey, it is evident that the

Figure 21. Distribution of A. torquatus costaricensis.

subspecies of A. torquatus generally fre-
quent the subtropical zone and quite regu-
larly reach the tropical zone. The one ex-
ception to this is A. t. assimilis which is
strictly a temperate zone form.

Habitat. — A. torquatus is a species of
thick second-growth and other dense vege-
tation in fairly humid areas. Slud (1964)
pointed out that in Costa Rica A. brun-
neinucha prefers heavy growth witliin tall
forest, and A. gutturalis [= aJbinucha] se-
lects bmsh in tlie open, while A. torquatus
occupies the bushy borders of forests, an
intermediate habitat. My observations of
these species in Mexico and Ecuador con-
firm this.

Habits.—Skutch (1954) has written a
brief life history account of this species in
Costa Rica.

A. torquatus frequently occurs alone or
in pairs, in contrast to the somewhat more
gregarious species such as A. rufinucha,
brunneinucha, or schistaceus, although at
times it does occur in moderate-size groups.

In Ecuador on several occasions, I have
observed the species foraging on the ground
and turning leaves with its bill. I have never

352 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

^^'

poliophry s

torquatus

•

f imbriatus

borelli

3'*

Figure 22. Southern races of A. torquatus.

seen it use its feet for this purpose. Skutch
(1954) also commented on this use of the
bill and noted that the bird feeds on small
invertebrates and at times eats decaying
leaves.

Its voice is thin and v/eak. In Ecuador, I
noted a song as zeep, which-a-whee, with
the tone dropping on the a and rising on the
ichee. Skutch (1954) notes that the song is
given from, or close to, the ground and de-
scribes the male's song as being squealy,
high-pitched, tuneless, and rapid; the fe-
male's song is similar but even weaker. I
noted two calls in Ecuador, a high, metallic,
zeep and a soft throaty chuck. Skutch
(1954) described the call as similar to that
given by a Cardinal {Cardinali.s cardinalis) .
Slud (1964:383) described the voice as an

insect-like trill or as the "tinkling of a fine
silver chain"; he also noted a cherrr which
must be what I described as chuck.

Breeding in Costa Rica is from February
to September ( Skutch, 1954 ) , in northern i
Venezuela it is from May to July ( Schiifer i
and Phelps, 1954). There is no information
from farther south. Skutch (1954) noted
that the species builds a bulky nest in dense
tangles from one to six meters above the
ground; two white or very pale blue eggs
are laid and incubated only by the female.

MorpJiological variation. — As has been
discussed above, morphological variation is
extensive within A. torquatu.s. The pectoral
band appears in ])a.siUcus\ perijanus, phaeo-
pletirus, phijiias. laren.sis, poliophrys, tor-
quatus, and f imbriatus (faint) and is absent

Biology and Evolution of Atlapetes • Paynter 353

in costaricensis, assimilis, nigrifrons, and
horelli (Fig. 20). Because of the irregular
distribution of this character, one is led to
suspect that the absence of the pectoral
band might be related to sympatry or
parapatry with A. brunneinucha, the species
that A. torquatus so closely resembles mor-
pliologically, as well as in behavior and
altitudinal distribution. However, no cor-
relation can be found.

The color of the superciliary is about
equally divided between races with gray
stripes and races with white stripes (Fig.
20). The northeastern and southern Andean
forms have white superciliaries; nigrifrom
of southwestern Ecuador and northwestern
Peru has a gray superciliary with white
lores; the remaining taxa have gray super-
ciliaries. This pattern does not seem to be
related to the presence or absence of a pec-
toral band, or to be correlated with contact
with other species of Atlapetes.

There is considerable, but not extreme,
variation in the color of the flanks and
undertail coverts (gray-green, brownish,
etc.) and the extent to which this color
suffuses the white belly. There are also
minor variations in the width and length of
the medial head stripe and in the color of
the back and tail. No patterns in these
variations are recognizable.

Males are slightly larger than females,
but I am unable to detect any significant
difference in size (wing, tail, culmen) be-
tween like sexes of the various races for
which I have adequate series of specimens
{costahcensis, basilicus, assimilis, and nigii-
frorvs); measurements of specimens in the
remaining races all fall within the ranges
] noted in the four large series. I conclude
that if there are racial differences in size
they are doubtless very small and certainly
of no biological significance.

ORIGIN AND INTERRELATIONSHIPS
OF THE TAXA

Introduction

It is now generally believed that the
exceptional abundance of Neotropical avian

species, as well as that of other animals and
plants, is mainly the result of Pleistocene
fluctuations in both temperature and pre-
cipitation. These climatic changes resulted
in the expansion and contraction of forests
and grasslands in the lowlands, as well as in
vertical shifts in biomes in the mountains.
These changes, in turn, led to the shrinkage
and even disappearance of some popula-
tions during one phase of the cycle while
during the alternate phase the populations
expanded their ranges. Species differenti-
ated to various degrees, disappeared, or re-
mained unaltered during the restrictive
phases while in the expansive stages they
kept their identity, were absorbed by other
populations, disappeared altogether, or even
further differentiated. Over the course of
several cycles of varying duration and in-
tensity, the opportunities for permutations
were enormous so that now, at best, only
broad patterns of speciation or those of re-
cent occurrence can be discerned.

This theory has been developed princi-
pally by Haff er ( for a summary see Haff er,
1974), who has applied it mainly to the
tropical lowland forest avifauna. He has
postulated 16 major areas in Central and
South America that served as forested refu-
gia for birds during the most arid periods
of the Pleistocene. Because cHmatic fluctu-
ations affected the mountains as well as the
lowlands, it was not surprising when in a
study of the Atlapetes schistaceus species-
group (Paynter, 1972) it became evident
that some subtropical /temperate zone taxa
seemed to have had their origin in close
proximity to the lowland refugia postulated
by Haffer. Furthermore, it was also ap-
parent that the origin of certain other taxa
could be explained only if there had existed
additional refugia in pockets on the slopes
of the mountains.

The present study completes the review
of the genus Atlapetes and refines and ex-
pands some of the zoogeographic concepts
developed earlier.

The 24 species that constitute Atlapetes
fall into four natural sulnmits that reflect

354 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

the evolutionary history of the genus. The
order in which these four subunits are
treated below is intended to show a gen-
eral trend from the most "typical" atlapetes
to those that approach other emberizine
genera. However, because a linear ar-
rangement sometimes does not permit all
related taxa to be near one another, and be-
cause e\'en the limits of emberizine genera
are fuzzy, only the grouping of the species
within the units should be considered as
significant.

Atlapetes albinucha superspecies

Two allospecies comprise this superspe-
cies, A. albinucha of Mexico, Central Amer-
ica, and Colombia and A. pallidinucha of
Colombia and Ecuador.

Atlapetes albinucha

A. albinucha has eight races (Figs. 3,
4). Two (coloratiis and azueren.sis) have
been described from Chiriqui and Veraguas,
western Panama, and are almost certainly
inseparable from brunnescens, also from
western Panama. Five of the remaining
races (griseipectus, fu.scipygius, parviros-
tris, brunnescens, and gutturalis) , ranging
from southwestern Mexico to Colombia, dif-
fer only slightly from one another, while
nominate albinucha of eastern Mexico is dis-
tinctly different, having a completely yel-
low venter rather than only a yellow throat.
Although distinctive, this is believed to rep-
resent a small genetic difference (see p.
330).

The sharp, albeit relatively minor, dis-
continuity in phenotype displayed by A. a.
albinuclui, in contrast to the very minor and
intergrading morphological variation in the
remaining races, even including A. a. gut-
turalis of Colombia which is isolated from
the Middle American populations by a siz-
able gap in eastern Panama, is provocative.
This suggests that A. a. albinucha may have
been isolated from the southwestern Mexico

and Central American populations for con-
siderable time, or at least more completely.
And, conversely, it could indicate that the
isolation of A. a. gutturalis may have been
more recent, or less complete.

Climatic changes in the Pleistocene might
account for this pattern. During the height
of a glacial period, when the vegetation
zones were lower, a population may have
been isolated in the lowlands of eastern
Mexico, possibly only north of the Isthmus
of Tehuantepec but more probably span-
ning the Isthmus of Tehuantepec to nortli-
eastern Chiapas as well; the latter distribu-
tion would explain the present occurrence
of morphologically similar populations on
either side of the Tehuantepec lowlands
(see Fig. 3). In isolation the northern
population either differentiated into distinc-
tive nominate A. albinucha or, more prob-
ably, simply did not change appreciably,
thereby retaining its similarity to A. pal-
lidinucha, its South American allospecies
(see p. 335).

On the other hand, during the Pleisto-
cene glaciations the low mountains of east-
ern Panama, which now lack subtropical
vegetation, except for a limited area on
Cerro Tacarcuna, and wliich are not now
occupied by this species, probably had a
suitable habitat. The presence of subtropi-
cal vegetation provide a series of stepping
stones facilitating exchange between the
Central America and Colombian popula-
tions. The populations of Central America
proper were at lower elevations than today
and probably were more nearly continguous
with one another than they are now. This
would account for their morphological simi-
larity which also suggests that their present
isolation is fairly recent. In addition, the
Colombian population doubtless was far-
ther west than now, owing to its presence on
the "discontinuous mountain bridge" ( Haf-
fer, 1974:15) which connected the Western
Andes and the mountains of Darien, across
what is now the lower Rio Atrato valley.

With the amelioration of the climate, the
low mountains of eastern Panama lost their

Biology and Evolution of Atlapetes • Paynter 355

subtropical vegetation and became unsuit-
able for the species. The Atrato "mountain
bridge" also disappeared and the species
retreated to the Andes. These events re-
sulted in a wide distributional breach.
Meanwhile, the subtropical zone moved
higher in the mountains of Chiapas. This
shift allowed the eastern and western popu-
lations to move closer together. This teeter-
totter effect would, therefore, bring the
northern isolate closer to the main body of
the species while at the same time it would
progressively increase the isolation of the
southernmost population. This seems to be
a reasonable explanation for the unusual
pattern of morphological variation now
seen.

Atlapetes pallidinvcha

A. paUidinucha generally resembles A. a.
albinucha. The principal difference is the
presence in A. paUidinucha of yellow lores,
an orange wash on the anterior part of the
central crown stripe, and, ventrally, faint
green striations, suggesting the plumage of
immature birds. The more southern forms
of A. albinucha differ more markedly in that
the vellow of the venter is confined to the
throat.

A. paUidinucha occurs in the Eastern and
Central Andes of Colombia and south to
central Ecuador, and just reaches Venezuela
(Fig. 5). In South America A. aUnnucha is
found on all three ranges of the Colombian
Andes, but in the Eastern Andes it is not
north of Cundinamarca (Fig. 4). (Neither
species has been recorded in the Eastern
Andes south of the vicinity of Bogota, but
this seems merely to be because of the ab-
sence of collectors; the southern portion of
the Eastern Andes are very poorly known;
see Fig. 1.) The two species have, there-
fore, overlapping ranges in the Central
Andes and in the midportion of the Eastern
Andes. They are, however, altitudinally
segregated with A. paUidinucha in the tem-
perate zone, or higher, and A. oWinucha in
the subtropics.

There are approximately 50 sites within
the zone of overlap where either one or the
other of the species has been recorded. At
only one site have both been reported. This
is at La Aguadita, Cundinamarca, at an al-
titude of about 2,000 m (Olivares, 1969).
This location is on the steep western slope of
the Eastern Andes. Altitudes exceeding
3,400 m are only a short distance from
La Aguadita. Presumably the two species
are segregated altitudinally within the re-
gion but were imprecisely cited as having
come from the same place.

Interrelationships

The morphological similarity and altitudi-
nal segregation of these two species sug-
gests that they are related but ecologically
incompatible. I consider them, therefore, to
be members of a superspecies. The re-
semblance of the Mexican race of A. al-
hinucha to A. paUidinucha of South Amer-
ica, or conversely the greater dissimilarity
between the two species where they are
altitudinally parapatric is suggestive. It
may indicate that nominate A. aUnnucha,
the Mexican race, is a little-changed off-
shoot from the same stem giving rise to A.
paUidinucha. The other races of A. al-
binucha, or at least the stock for these, may
have arisen while in proximity to A. paJ-
Udinucha and, therefore, diverged more
than did nominate A. albinucha which was
far to the north.

I have postulated above (p. 354) that the
race of A. aUnnucha in Colombia (A. a.
gutturaUs), although well-isolated from the
Central American populations of the spe-
cies, is morphologically similar to these pop-
ulations because this isolation is relatively
recent. On the other hand, the Mexican
race, A. a. aUnnucha. which is narrowly iso-
lated from another population, is morpho-
logically distinct because it is an older iso-
late that only recentK' has come in near
contact with another population.

The postulated origin of the races of A.

356 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

alhinucha and the origin of tlie allospecies
A. alhinucha and A. pallidinucha can be
reconciled as follows. Presumably the spe-
cies had their origin during a dry period in
the Quaternary when an ancestral popula-
tion became dixided between two (or
more) forest refugia. Probably A. pallidi-
nucha formed in a refuge at the head of the
Central and W^estern Andes ( the Caribbean
Colombian or Nechi Refuge of Haffer,
1974). The other allospecies, A. alhinucha..
may have been formed in isolation in a
refuge on the Pacific side of the Western
Andes (Pacific Colombian, or Choco, Ref-
uge) or, even more probable, in Central
America in the Caribbean Central American
or in the Caribbean Costa Rican Refuge ( all
refuge names from Haffer, 1974), or pos-
sibly in a refuge in Mexico north of the
Isthmus of Tehuantepec. Later with the
amelioration of the climate the two forms
moved out from their centers of origin and
came into contact. They had not diverged
much from one another, either morphologi-
cally or ecologically, but they were suffi-
ciently differentiated so that they were re-
productively isolated and retained their
identity. There may have been parapatry,
or partial sympatry, or, more likely, some
altitudinal overlap with A. pallidinucha the
higher taxon. In any case this may have re-
sulted in further divergence between the
two similar species. A. pallidinucha may
have been prevented from ranging into
Central America because of the lack of
temperate forest on the low mountains.
During the next phase of the climate cycle
A. pallidinucha may have withdrawn to the
Nechi Refuge, or even to the Napo Refuge
of eastern Ecuador, and another population
was isolated in the Eastern Cordillera of
Colombia; here the two existing races
differentiated. Meanwhile, A. alhinucha
pulled back to two or more refugia, the
northern one (probably in Mexico) holding
the population which had been farthest
from the zone of contact with A. pal-
lidinucha and a southern refuge in Central
America or northwestern Colombia, with a

population which had been in contact with
A. pallidinucha. This isolation allowed A.
alhinucha in its southern refuge to consoli-
date morphological and ecological diver-
gencies brought about because of com-
petition with A. pallidinucha, while the
population in the northern refuge remained
relatively stable in appearance, having been
far away from the zone of contact with A.
pallidinucha. During the next warm-moist
period (the current one?) the population
in the northern refuge (A. a. alhinucha)
moved higher in the mountains of south-
eastern Mexico, but was unable to cross
the low Rio Crijalva valley in Chiapas. The
population in the Central American refuge
spread throughout much of Central Amer-
ica and across into Colombia, later abandon-
ing eastern Panama as the climate warmed
and subtropical vegetation disappeared.
Because of its attenuated range, clinal mor-
phological changes have developed, result-
ing in the weakly differentiated races now
recognized.

Although A. pallidinucha papallacta is re-
markably similar to A. nifinucha haroni
(see p. 334), I do not believe they are par-
ticularly closely related.

Atlapetes rufinucha species-group

The 11 species within this group are
presumed to have had a common origin,
althcnigh all do not seem to have split off
at the same time. With only one or two
exceptions, and these are based on doubtful
records, none of the species is sympatric
with any other member of the species-
group, which, of course, lends credence to
the belief that this is a natural assemblage
of closely related taxa.

Atlapetes rufinucha superspecies

This superspecies is composed of A.
rufinucha, a widespread polytypic species,
and A. melanocephalus, a monotypic ende-
mic of the Sierra Nevada de Santa Marta,
the isolated massif in northern Colombia.

Biology and Evolution of Atlapetes • Fayntcr 357

Atlapetes rufinucha

A. rufinucha is the most widespread of
the 11 species within the species-group,
occurring from westernmost Venezuela to
BoHvia. There are a number of gaps in its
range (Figs. 6, 7). While it is present in
the Sierra de Peri j a, a northward extension
of the Eastern Andes on the Venezuela-
Colombia border, it has not been found in
the Eastern Andes except for a few dubious
records from "Bogota" (A. r. simplex, known
only from native "Bogota" specimens). It
does not occur on the eastern slope of the
Central Andes, but is found at the northern
tip of this range, as well as on its western
slope at the head of the Cauca valley. In
the \\^estern Andes, it is found only on the
eastern slope, also far up the Cauca valley.

In Ecuador it is distributed on both
slopes, but is absent for about 250 km in
the central portion of the eastern slope.
The species ranges a short distance down
the western slope of northern Peru and in
the east reaches central Peru. There is then
a gap before the species recurs in southern
Peru and eastern Bolivia.

Its ecological requirements are broad. It
occurs in the subtropics, sometimes in the
temperate zone, and in dry to moderately
moist vegetation. Its altitudinal range is
also notably extensive, covering about 3,000
m. In general this is a common and abun-
dant bird, as one would expect from its
tolerance of a wide range of ecological con-
ditions. Unfortunately, the versatility of the
species makes it impossible to speculate on
its place of origin. The gaps in its distribu-
tion, however, are valuable clues in recon-
structing the history of other species in this
species-group (see A. tricolor superspecies,
A. albofrenatus, and A. leucopis).

Atlapetes melanocephalvs

A. melanocephalus is the second allospe-
cies of the rufinucha superspecies. It is one
of the two atlapetes in the isolated Sierra
Nevada de Santa Marta (Fig. 8) and is
closely allied to A. rufinucha and might

even be considered a particularly well-
marked race of that species. It differs from
A. rufinucha in having a fully black head
(not chestnut), a black chin and upper
throat, and silvery ear coverts. These char-
acters are found in varying degrees in some
races of A. rufinucha (see Table 1). For
example, the black of the chin and throat
occurs in A. r. mehinolaemus. Also, in A. r.
phelpsi, the race nearest to Santa Marta,
there is a broad black band on the forehead
and along the sides of the head which seems
to be a step toward a fully black head; its
ear coverts are nearly as silvery as in A.
melanocephahis.

Interrelationships

There seems little doubt that A. melano-
cephahis had its origin in A. rufinucha, and
probably from stock from which arose
A. ;•. phelpsi. It is probably the youngest
species within the A. rufinucha species-
group.

A. melanocephalus occurs from the up-
per tropical zone ( 600 m ) through the sub-
tropical zone ( 2,400 m ) . It is the only mem-
ber of the species -group to occur so low.
Although this altitudinal range coincides
with that of A. torquatus on Santa Marta,
A. melanocephalus, if it is like A. rufinucha,
is probably more arboreal and, therefore, is
not ecologically competitive with it. There
are no additional atlapetes on Santa Marta
and presumably this allows A. melanocepha-
lus a wider range in altitude than is usual
in the genus.

Atlapetes tricolor superspecies

The three allospecies of this superspecies
are morphologically very similar, the princi-
pal difference being in the color of the
crown, which in A. tricolor is gold, in A.
flaviceps is yellow, and in A. fuscoolivaceus
is blackish. Immature A. tricolor has a dull
crown, resembling that of adult A. fuscooli-
vaceus-, no immature examples of A. flavi-
ceps are known. Interestingly, immature A.
rufinucha bears a strong resemblance to A.

358 Bulletin Miiseiini of Comparative Zoology, Vol. 148, No. 7

tricolor, suggesting a relationship between
the two taxa. The distribution of the two
species ( see below ) further strengthens this
behef.

Atlapetes tricolor

A. tricolor has an extended range (Fig.
9), beginning on the western slope (and
one doubtful record for the eastern slope)
of the \\'estern Andes of Colombia and
extending down the western slope of Ecua-
dor, but becoming very sparse in the south,
presumably because of increasing aridity.
It reappears on the eastern slope in central
Peru, after a gap of about 1,000 km. It oc-
ciu-s from the humid upper tropical zone to
the upper subtropical zone in the north and
in the subtropical and temperate zone in
Peru.

Atlapetes fuscoolivaceus and
a. flaviceps

Both A. fiiscoolivoceus and A. flaviceps
have very restricted ranges on the eastern
slope of the Central Andes (Fig. 9). A.
flaviceps occurs in a deep valley on the
south slope of Nevada de Tolima and A.
fuscoolivaceus at the headwaters of the Rio
Magdalena in a cul de sac where the East-
ern Andes swing abruptly west to join the
Central Andes. Both species are known
only from the subtropical zone. There has
been little ornithological work on the east
side of the Central Andes; it is possible that
the ranges of these two species may be
more extensive than now known.

Interrelationships

A. flaviceps and A. fuscoolivaceus appear
to be remanent offshoots of the stock that
produced A. tricolor. Presumably ancestral
A. tricolor was once confined to the Pacific
Colombian (or Choco) Refuge, as defined
by Haffer (1974), and later expanded its
range down the western slope of Ecuador,
crossing over to the east at the low passes
in southern Ecuador and northern Peru. A

subunit of the same ancestral stock seems
also to have been restricted to the eastern
side of the Central Andes, perhaps in the
Caribbean Colombian (Nechi) Refuge.
During a warm-moist period, it may have
spread up the Magdalena valley along the
eastern slopes of the Central Andes, only to
retreat later to one area at the head of the
valley, where A. fuscoolivaceus is now
found, and to another restricted region 300
kilometers to the north, where A. flaviceps
occurs.

The reason why A. fuscoolivaceus and A.
flaviceps are restricted to two small areas
doubtless is related to the wetness of the
subtropical habitat at these points. Both
places are at the head of valleys which are
cut deep into the Andes. Presumably these
regions receive much more precipitation
than points that are farther east, away
from the mountains, and nearer the arid
upper Magdalena valley. Thus both species
are probably unable to move down the val-
leys or up over their sides because of un-
suitable habitats. These two places are
certainly not the only sites with moist sub-
tropical vegetation along the entire eastern
slope of the range. Future work may re-
veal additional populations in pockets at
the heads of other deep valleys.

Geographical Displacement

The wide geographical breach between
the subspecies of A. tricolor is puzzling, as
is the absence of the species on the eastern
slope of Ecuador. Displacement by another
species would seem an explanation. A.
rufinucha, which in immature plumage is
similar to A. tricolor, appears a likely spe-
cies. The races A. r. haroni and latinuchus
more or less fill the breach on the eastern
slope from southeastern Ecuador to central
western Peru (Fig. 6). A. t. tricolor then
occurs from central to southern Peru, and
this is followed by two races of A. rufinucha
ranging from southern Peru to southeastern
Bolivia ( Figs. 7, 23 ) . Although this would
appear to be a classical example of a
geographical replacement, there are some

Biology and Evolution of Atlapetes • Paynter 359

flaws, the most important being tliat in
general the species are found at different
altitudes. In western Colombia and Ecua-
dor, the two species have generally overlap-
ping ranges, but A. tricolor is invariably at
lower elevations. If the two species are
competitors, one would expect that in the
absence of one, the other might expand its
altitudinal range. However, A. rufimicha
in eastern Ecuador, where A. tricolor is ab-
sent, has the same vertical range as it does
in western Ecuador, where A. tricolor is
found. In Peru A. f. tricolor is at moderately
higher elevations than is A. t. crasstis in
Colombia and Ecuador and, therefore, it
occupies the lower part of the altitudinal
range of A. nifinucha. In other words, A.
tricolor seems to have expanded slightly up-
ward in the absence of A. rufimicha. I can
find no evidence that A. rufimicha drops to
lower altitudes in Peru where A. tricolor is
lacking; it does, however, occur as low as
600 m in Bolivia. Unfortunately, there are
available only about 100 records for both
species in a range covering over half the
length of the Andes. While broad outlines
of altitudinal preferences are obtained, it is
hardly likely that more subtle differences in
altitude between species and races will be
revealed by these few data.

The manner in which A. tricolor replaces
A. rufimicha and the resemblance of imma-
ture A. rufimicha to A. tricolor lead to the
conclusion that the two are members of the
same species-group. It follows that if A.
tricolor is a member of the species-group
then its allospecies, A. flaviceps and A.
fuscoolivaceus\ must also be in the group.
The presence of these two species on the
eastern slope of the Central Andes, occupy-
ing a gap in the range of A. rufimicha, fur-
ther reinforces the belief that these are
indeed members of a closely related as-
semblage.

Atlapetes albofrenatus

A. a. albofrenatus is one of the most dis-
tinctive taxa of the A. rufimicha species-
group by virtue of its white throat, heavy

malar streaks, and green back; however,
in the race A. a. meridae, the white throat
and heavy streaks are reduced, and the bird
is much less distinctly different from A.
rufimicha. In any case, these characters are
minor. The placement of A. albofrenatus in
association with A. rufimicha is reinforced
by two points. First, this is an active bird
whose behavior is similar to that of A.
rufimicha in contrast to the slower, more
secretive movements of most atlapetes. Sec-
ond, and more important, the species occurs
within a breach in the range of A. rufinucha
in the Eastern Andes from Bogota north to
southwestern Venezuela (Fig. 23). Of par-
ticular note is the fact that it does not occur
in the Sierra de Perija, the northward pro-
jecting spur of the Andes which is occupied
by isolated A. r. phelpsi, but it does cross
the barrier created by the depression
formed by the Rio Torbes, which separates
the Eastern Andes from the Andes of
Merida. The only possible instance of sym-
patry is in the vicinity of Bogota where A. r.
simplex is said to occur, but this race is
known only from several native "Bogota"
specimens and certainly comes from else-
where, probably in the little-known Eastern
Andes south of Bogota. The altitudinal
range of A. albofrenatus is somewhat more
restricted than that of A. rufinucha, but the
species displays the same versatility in habi-
tat preference.

Thus there is no doubt that A. albofrena-
tus is related to the A. rufinucha species-
group. Its relative distinctiveness indicates
it is not part of the A. rufimicha species
branch, but was derived from the same
stock that produced A. rufinucha. It would
seem to have originated in the Eastern An-
des, simply because that is where it is now
found. No Pleistocene refuge for subtropi-
cal forest forms has been proposed in the
area now occupied by A. albofrenatus, al-
though Haff er ( 1974 ) believes one existed
farther north (the "Catatumbo Refuge").
The western slopes of the Sierra Nevada
del Cocuy, the highest peak in the Eastern
Andes, may have served as refugium. The

360 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

Figure 23. Geographic replacement witiiin the A. rufinucha species-group in South America. Areas of doubtful
sympatry are marked "?".

Biology and Evolution of Atlapetes • Paynier 361

mountain is at the head of the long and arid
Cliicaniocha valley running west to the
-\higdalena and may have caught moisture
during dry periods and served as a refuge
for forest inliabitants, even if the valley it-
self was arid. This is the region occupied
by nominate A. alhofrenatus. On the other
hand, the fact that A. alI)ofrenatiis meridae,
the race of the Merida Andes, is less dis-
tinct from A. rufimicha than is A. a. alho-
frenatus, suggests that the Merida Andes
may ha\'e served as the initial isolation area
and that the species later crossed the Rio
Torbes barrier where it further differenti-
ated. Both hypotheses could be embel-
lished, but there seems no way to establish
which area could have served as a specia-
tion center. Indeed, A. alhofrenatus may be
a relict and not autochthonous to either
place.

Atlapetes leucopis

As I have observed before (Paynter,
1970), A. leucopis resembles A. rufimicha
melanolaemus of Peru, the most distinctive
race of A. rufimicha, except for its much
larger size, white eye ring and eye stripe,
and green underparts.

It seems to be sympatric with A. rufi-
nucha at Palmas, Azuay, on the eastern
slope of Ecuador, but does not coexist at
the other two localities from which it is
known, viz. La Plata, Colombia, which is
on the east side of the Central Andes and
Cen-o Pax, on the east slope in southern
Colombia (Fig. 23). The Palmas locality
is suspect, however. The single bird from
there was obtained by M. Olalla (Berlioz,
1932), a member of the family of profes-
sional collectors that is noted for inaccu-
rately labeled specimens. From a few hours
spent at Palmas, I know, that A. rufimicha,
which Olalla also records from here, does
indeed occur at this locality; I believe that
A. leucopis may be somewhere in the gen-
eral region but not sympatric with A. rufi-
nucha.

The two Colombian sites for A. leucopis
are to the east of the range of A. rufinucha

and the Ecuadorian locality is at the north-
ern edge of the range of A. rufinucha, which
reappears again about 250 kilometers far-
ther north in Ecuador. Thus it seems that
A. leucopis is a geographical replacement
for A. rufimicha, although it is possible that
allopatry may break down in Azuay at the
southern end of the range of A. leucopis and
the northern end of the range of A. rufi-
nucha.

A. leucopis is not known to be sympatric
with any other member of A. rufimicha
species-group, but it should be noted that
A. fuscoolivaceus occurs in Colombia at
Moscopan (alt. 2,400 m), Huila, which is
only .32 km west of La Plata (alt. 2,350 m),
and in the same valley.

It would seem that A. leucopis bears a re-
lationship to A. rufinucha similar to that
existing between A. rufinucha and A. alho-
frenatus, i.e., A. leucopis appears to have
arisen from the same ancestral stock as A.
rufinucha, but it is not a derivative of A.
rufimicha. It is not possible to speculate
on its place of origin.

Atlapetes pileatus

A. pileatus, the Mexican endemic (Fig.
12) is a small pallid version of A. rufinucha.
It is isolated from the remainder of the
species-group by all of Central America. It
appears to be a relict that originated in
Mexico or Central America, but it is not
possible to speculate further.

Atlapetes fulviceps superspecies

There are three allospecies within the
A. fulviceps superspecies, viz. A. fulviceps,
A. personatiis, and A. semirufus. All are
morphologically very similar inhabitants of
the subtropical zone, and all ha\'e a marked
resemblance to A. rufinucha, but differ from
A. rufimicha in that the black of the sides of
the head is replaced by chestnut and, in
some taxa, the chestnut extends to the throat
and breast. A. fulviceps. which is monotypic,
is found in Bolivia and northwestern Argen-
tina ( Fig. 14 ) . A. personatiis, with six well-

362 Bulletin Museum of Comparative Zoology, Vol 148, No. 7

marked races, occurs in the "pantepuis" of
southwestern Venezuela (Fig. 13). Both
species are at or near the outer edges of the
distril)ution of the genus. A. persomitus is
the sole atlapetes in its region; A. rufinucha
occurs in the general area with A. fulviceps,
in the latter's northern range, but has been
recorded as svnipatric with it only at
Tilotila (alt. 2,150 m). La Paz, Bolivia. The
Tilotila records are those of Buckley, whose
collections were made from 900 to 3,700 m
(Sclater and Salvin, 1879). The absence of
further evidence of sympatry suggests that
Buckley's specimens were altitudinally sep-
arated, but which were at the higher eleva-
tion is unknown because the altitudinal
records from elsewhere overlap.

Highly polytypic A. seminifus is the spe-
cies of the Eastern Andes of Colombia and
the coastal mountains of Venezuela ( Fig.
13). It may be sympatric with A. albofrena-
tus near Bogota, but the data suggest that
A. seminifus occurs at higher altitudes.

Presumably A. persomitus was derived
from early A. seminifus. Mayr and Phelps
(1967), apparently following Chapman
(1931), state that A. persomitus is most
closely related to far-distant A. fulviceps.
However, morphological evidence is incon-
clusive and derivation of A. personatus from
A. seminifus seems more logical, simply be-
cause they are geographically closer.

The long gap between A. seminifus of the
northern Andes and A. fulviceps of the
southern Andes is provocative and suggests
that the two allospecies are separated by
another species or group. The A. rufimicha
species-group fills the breach without any
indication of sympatry, except for the very
dubious record of A. rufinucha simplex
from native "Bogota" specimens (Fig. 23).
This close fit does not seem to be an artifact.
A. fulviceps superspecies not only is pheno-
typically similar to the A. rufimicha species-
group but must also be very closely related
and, hence, unable to coexist with it.

Although (me could argue well for the
inclusion of A. persomitus, A. seminifus, and
A. fulviceps within the A. rufinucha species-

group, there is a morphological cohesiveness !
among the three species which probably
indicates that this group branched from
the ancestral stem stock earlier than the
indi\'idual species making up the A. rufi-
nucha species-group.

Atlapetes citrinellus

A. citrinellus, of Argentina, is the south-
ernmost representative of the genus (Fig.
15 ) . It stands apart from other members of
the genus and obviously is an aberrant rep-
resentative. It does bear some resemblance
to A. fulviceps without the chestnut crown
and cheeks. The two may have had an an-
cestral branch in common.

Atlapetes schistaceus species-group

This group of six closely-related species
is distributed through the Andes from west-
ern Venezuela to southern Peru (Fig. 24).
The species replace one another geographi-
callv in a manner reminiscent of the mosaic
pattern of the A. rufinucha species-group.
The group was analyzed earlier ( Paynter,
1972) and only brief outlines are gi\'en
here.

Atlapetes schistaceus

This species has the widest distribution
of any species within the A. schistaceus
species-group, ranging disjunctly from the
Cordillera de Merida, Venezuela through
the three ranges of the Colombian Andes
and thence on the eastern slope to central
Ecuador. The species reappears in central
eastern Peru and then again in southeastern
Peru (Fig. 24). It is mainly a humid tem-
perate region form but does reach the sub-
tropical zone on occasion. Its total range is
from 1,850 to 3,750 m. Color and pattern
vary geographically; 12 races have been de-
scribed. It is, therefore, morphologically
the most variable of all atlapetes.

Altapetes nationi

A. nationi (Fig. 24) is the southern An-
des western slope counterpart of A. schista-

Biology and Evolution of Atlapetes • Faynter 363

\ ■

\ ^

V A

schistaceus

•

n a ti o n i

leucopterus

albiceps

pallidiceps

O
■
▲

ruf igenis

Figure 24. Distribution of the A. schistaceus species-group.

364 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

ceus, occurring from soutliwesteni Ecuador
south to Arequipa, Peru, with ahnost the
same altitudinal range as A. schistaceus but
apparently with a greater tolerance for drier
habitats. It is the only Athipetes to range
so far south on the western side of the Per-
uvian Andes. It also has a fragmented dis-
tribution and is moiphologically varied,
with eight races being recognized.

Atlapetes leucopterus

This, the smallest bird in the genus, is
found on the western slopes from north-
western Ecuador to northern Peru, and re-
cently was collected by J. W. Fitzpatrick
on the eastern side of the Andes in the
Cordillera del Condor, Cajamarca, Peru
( Fig. 24 ) . It has a notably wide altitudinal
range, having been found from 600 to 2,900
m, and appears to be common in rather
xerophytic areas, but it also occurs in moist
regions.

Distributional records show three clusters,
viz. northern Ecuador, central Ecuador,
and southern Ecuador and northwestern
Peru (Fig. 24). The northern and central
Ecuador populations represent one subspe-
cies and the southern Ecuador and north-
western Peru populations another race.
The recently discovered population in
brushy edges of cloud forest in the Cordil-
lera del Condor, to the east of the pre-
viously known Peruvian range, represents a
very distinctive form that has not yet been
named.

A. leucopterus is sympatric with A. nfl-
tioni and A. albiceps in the dry, low moun-
tains of southern Ecuador/northern Peru.
These are the only known instances of sym-
patry within the species-group. The area
of sympatry is in a region that probably has
been particularly sensitive to variations in
climate, owing to its low mountains and
proximity to the arid Pacific coast. The
vegetation has doubtless varied greatly.
The instability would have made the region
unsuitable for the long-term presence of
any species of Atlapetes. The species, now
meeting there and overlapping, seem to

have had their origins elsewhere — tmtioni,
and probably albiceps, to the south and
leucopterus to the north (see Paynter, 1972:

317-318).

Atlapetes albiceps

A. albiceps, a monotypic taxon, is an-
other western slope form. It is found from
southernmost Ecuador to Cajamarca, north-
ern Peru ( Fig. 24 ) . It is an arid zone spe-
cies with an altitudinal range from 250 to
1,500 m.

Atlapetes pallidiceps

This is another arid area form, ranging
from about 1,500 to 2,100 m. It is endemic
to the valley of the upper Rio Jubones and
its tributaries, southwestern Ecuador (Fig.

24).

Atlapetes rufigenis

A. rufigenis is a large species with a pop-
ulation in the drainage system of the Rio
Marafion, northern Peru and another, ra-
cially distinct, population far to the south
in the Rio Apurimac region (Fig. 24). It
occurs from 2,750 to 4,000 m and seems to
frequent mesic underbrush.

Atlapetes torquatus species-group

There are four species within this species-
group. The presence of a chestnut pileum
in A. brunneinucha sharply distinguishes it
from A. torquatus superspecies, but except
for this one character, there is little morpho-
logical difference. All four species are surely
derived from a common stem. However,
because A. brunneinucha is frequently sym-
patric with A. torquatus, the two must be
old, well-established, species, in contrast to
the species comprising the A. rufinucha
species-group which, because they seem
unable to coexist and therefore form a com-
plex geographical mosaic, are presumably
actively speciating.

Atlapetes brunneinucha

A. brunneinucha has the most extensive
range of any atlapetes (Figs. 16, 17) but

19::),

Biology and Evolution of Atlapetes • Paynter 365

shows remarkably few morphological varia-
tions, except for the absence of a breast
band in the races apterus, aUinornatus, and
inonuitus, three very small populations.
This is a notable contrast to the variability
within A. torqiiatiis superspecies, which is
almost as widely distributed.

The lack of morphological variability is
probably because A. hrunncimicha inhabits
the interior of moist subtropical forest. This
habitat doubtless continued to exist in
abundance and with few discontinuities
even during the peak of climatic deteriora-
tion when lowland forests withdrew to iso-
lated or semi-isolated refugia. Indeed, with
the lowering of vegetation zones on the
mountains, subtropical forest may have be-
come more abundant than at any period,
owing to its presence on the vast shoulders
of the mountains.

The morphological differentiation that
has occurred in the races apterus, aUinorna-
tus, and inornatus is doubtless the result of
isolation, but why the breast band is the
variable character is unknown. A. b. ap-
terus is on an isolated mountain standing in
lowland forest in Veracruz; A. b. aUinorna-
tus is on a northward-projecting spur off the
main range of the Venezuelan Andes. There
is no geographical feature associated with
the range of A. b. inornatus, but its origin
may be attributed to isolation brought abtnit
by climatic changes. Note that the arid
coastal belt of western South America
reaches northern limits at about the same
latitude as the range of inornatus. During a
warm-dry period the arid zone must have
extended higher on the western slopes of the
Andes. A. b. inornatus may have originated
in a surviving pocket of wet, semitropical
forest high on these slopes. A refugium in
the same area has already been proposed to
explain the origin of A. leucopterus (Payn-
ter, 1972:317).

Atlapetes torquatus superspecies

A. virenticeps, the northernmost repre-
sentative of this superspecies (Fig. 18), is
a relict population. Its plumage is reminis-

cent of immature A. torquatus, but whether
this indicates that the plumage of virenti-
ceps represents the ancestral pattern of the
superspecies, or is a case of reversion from a
more "adult" plumage, or is an entirely
"new" plumage is unanswerable.

The origin of A. atricapiUus (Fig. 19) is
perhaps one of the most intriguing but baf-
fling problems of this nature in the genus.
If, as it seems now, A. atricapiUus and A.
torquatus are parapatric, the former is prob-
ably a fairly recent offshoot. The fact that
it is a lower altitude species than A. torqua-
tus, and also occurs mainly in the northern
extremities of the Andes suggests that it
may have originated on outliers of the main
Andean ranges, but more specific specula-
tion is not possible until we have a better
knowledge of the range of the species.

The morphological variability of A. tor-
quatus (Figs. 19-22) stands in contrast to
the uniformity of its sister species A. brun-
neinucha. The difference may lie in their
habitats. While both are subtropical forms,
although A. torquatus also ranges higher, A.
torquatus is an edge dweller, often frequent-
ing second-growth. Under natural condi-
tions, edges and second-growth are gen-
erally uncommon and scattered, which
means that populations of A. torquatus
probably never have been as large or as
contiguous as those of A. brunneinucha, and
presumably this would have resulted in in-
creased polymorphism.

A. torquatus seems to be somewhat less
restricted to a given biome than many
atlapetes, which may be indicative of its
nature as a generalist and exploiter of tem-
porary changes in habitats. However, of
particular interest is the race A. t. assimiUs
which is strictly a temperate zone form. It
is also the only race within the range of the
allospecies A. atricapiUus. This is strongly
suggestive of altitudinal displacement, with
A. t. cissimiUs moving higher to accommo-
date A. a. atricapiUus in the upper tropical
zone. However, it should be borne in mind
that assimiUs has an extensive range, only a
small part of which is known to be near that

366 Bulletin Museum of Comparative Zoology, Vol. 148, No. 7

of A. atricapillus, and that its restriction to
the temperate zone may ])e for other rea-
sons.

CONCLUSIONS

The species of Atlapete.s are most abun-
dant at middle elevations in the northern
Andes, and it is assumed that the genus
had its origin in that region. This would
mean that the genus can be no older than
the uppermost Pliocene/lowermost Pleisto-
cene, because this was when the main up-
lift of the Andes took place (Haffer, 1974:
130). It could, of course, be considerably
younger.

The majority of the species seem to have
arisen in the northern Andes, but five of the
24 species (pileatiis, fulviceps, personatus,
citrineUus, and virenticeps ) must have origi-
nated well outside of this area and four
species ( tuitioni, albiceps, nifigenis, and
paUidiceps) may have arisen in tlie mid-
portion of the Andes.

The present pattern of distribution, as
well as the pattern of speciation, show many
indications of having been greatly influ-
enced by climatic changes. The connection
between climate and speciation is some-
times seen directly, as when the birds have
ranges that coincide with patches of vege-
tation and tlie patchines could only have
developed through the disappearance of
suitable intervening areas. The fragmenta-
tion of belts of vegetation must have been
caused by climatic changes. For example,
A. flaviceps and A. fiiscoolivoceus seem to
have arisen when populations of ancestral
A. tricolor were stranded in isolated pockets
of moist subtropical forest on the eastern
slopes of the Central Andes of Colombia,
above the arid upper Magdalena valley.
Another example may be found in A. pal-
lidiceps, a distinctive form isolated in the
arid valley of the upper Rio Jubones, east-
ern Ecuador. A. pidlidiceps is a derivative
of A. Ieiicoptertis\ a species of generally
wetter areas. In this case the population
was left behind by a shrinking belt of moist
forest but instead of finding refuge in a

wetter pocket (there are none in the region)
adapted to a drier situation.

Other indications of the effect of climatic
changes on speciation may be seen in the
patterns of distribution. For example, the
failure of A. albinucha gutturalis of Colom-
bia to diverge appreciably from Central
American populations, although well-iso-
lated, while nominate A. albinucha of Mex-
ico is markedly different from a series of
Central American populations, even though
less than 100 kilometers apart (see p. 329),
can only be explained by a shifting of
biomes through climatic changes. Another
illustration is found in the intricate distri-
butional mosaic of the A. rufinucha species-
group (Fig. 23) which certainly must have
arisen through a series of climatic oscilla-
tions that caused a series of contractions and
expansions in several different populations.
There seems no other explanation, for ex-
ample, for the leap-frog pattern displayed
by A. tricolor and A. rufinucha or for the
appearance of populations of A. albofrena-
tus between those of A. semirufus.

The 24 species of Atlapetes cluster into
four main groups which seem to represent
four multi-branched evolutionary lines.
These pathways and the interrelations of
the component species are shown diagram-
matically in Figure 25. Starting at the bot-
tom of the diagram and reading clockwise,
the following information is indicated.

A. brunneinucha and A. torquatus (with
the three allospecies, atricapiUus, torquatus,
and virenticeps) share a common origin
and fonn one evolutionary line, the A,
torquatus species-group. Superspecies A.
albinucha ( with allospecies albinucha and
palUdinucha) is a second main branch. The
third, and largest, branch is made up of 12
species. A. pileatus, A. leucopis, A. albo-
frenatus, A. rufinucha superspecies (with
allospecies rufinucha and melanocephalus),
and A. tricolor superspecies (with allospe-
cies flaviceps, tricolor, and fuscoolivaceus)
form the core of this branch while A. fulvi-
ceps superspecies (with allospecies fulvi-
ceps, semirufus, and personatus) is an off-

Biology and Evolution of Atlapetes ' Paynter 367

alb

Figure 25. Interrelationships within the four main evolutionary paths in the genus Atlapetes. The relative
lengths of the lines and their angles are of no significance. See text (p. 366) for details.

shoot of the main branch, and A. citrinellm
is presumed to be a derivative of this off-
shoot. The fourth line of descent is the
A. schistaceus species-group. A. rufigenis
seems to have been an early offshoot, or it
^ may merely be distinctive because of its
distance from the center of the species'
distribution. A. albiceps and nationi, while
sharing a common origin from the main

branch, are too distinct to be considered
allospecies. A. schistaceus seems to repre-
sent the main line of e\'olution for this
unit. A. pallidiceps and A. leucopteriis are
another pair of species that come off a com-
mon point but also are too dixergent to be
treated as members of a superspecies.

Of particular interest are the A. rufinucha
species-group and A. schistaceus species-

368 Bulletin Muscutu of Comparative Zoology, Vol 148, No. 7

group, which form the two main exokition-
ary lines within the genus. The former
has 11 species (12 if citrineUiis is included)
falling into three superspecies, plus three
(or four) separate species, while the A.
schistaceiis species-group is composed of
six species, none of which is a member of a
superspecies. The species within each spe-
cies-group dYv distributed in an intricate
checkerboard pattern. Sometimes the spe-
cies are geographically separated, at other
times they are contiguous, and only \'ery
rarely are they partially sympatric. In other
words, the species within each species-
group, e\en those which are not allospecies,
almost invariably do ntjt have overlapping
ranges. This general allopatry implies that
the species are incompatible and seems to
indicate that they have only recently speci-
ated.

LITERATURE CITED

Bangs, O. 1908. Notes on birds from western

Colombia. Proc. Biol. Soc. Washington, 21:

157-162.
Berlioz, J. 1932. Nouvelle contrilnition a I'etnde

des oisean.x de I'Ecuador. Bnll. Mus. Nat. Hist.

Nat. Paris, 4:620-628.
Bertoxi, a. de W. 1925. Notas sobre aves del

Paragnay, Hornero, 3:279.
Blake, E. R. 1956. A collection of Panamanian

nests and eggs. Condor, 58:386-.388.
. 1958. Birds of Volcan de Chiriqni, Pan-
ama. Fieldiana: Zool. [Chicago], 36:499-

577.
Carriker, M. a., Jr. 1910. An annotated list of

the birds of Costa Rica including Cocos

Island. Ann. Carnegie Mus., 6:314-915.
Chapman, F. M. 1914. Diagnoses of apparently

new Colombian birds, II. Bull. Amer. Mus.

Nat. Hist., 33:167-192.
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us ISSN 0027-4100

Bulletin OF THE

Museum of

Comparative

Zoology

Natural History of Cerion VIM: Little

Bahama Bank-A Revision Based on

Genetics, Morphometries, and

Geographic Distribution

STEPHEN JAY GOULD AND DAVID S. WOODRUFF

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 8
8 SEPTEMBER 1978

NATURAL HISTORY OF CERION VIII: LITTLE BAHAMA BANK—
A REVISION BASED ON GENETICS, MORPHOMETRICS, AND
GEOGRAPHIC DISTRIBUTION'

STEPHEN JAY GOULD^
DAVID S. WOODRUFF^

Abstract. Close to a dozen names are now
available to describe variation in Cerion on the
islands of Little Bahama Bank. These names,
plotted in the supposed areas of their occurrence,
form the "crazy-quilt" distribution pattern, tra-
ditionally, associated with Cerion and ascribed to
haphazard transport by hurricanes. We, on the
other hand, find remarkably stable patterns in
Cerion throughout the northern Bahamas. More
than 200 "species" can be reduced to a single,
unerringly predictable distribution and interaction
of two imperfectly separated entities: a ribby
morphotype associated with coasts that abut the
edges of the Pleistocene banks, and a mottled
morphotype from interior areas and coasts adjacent
to bank interiors. We find the same distribution
throughout Little Bahama Bank and reduce the
current taxonomy to two semispecies: C bendalli
Pilsbry and Vanatta ( the mottled morphot>pe ) and
C ahacoense Pilsbry and Vanatta (the ribby
morphotype ) . The distribution of these semispecies
maps the edges and interiors of Pleistocene banks,
as described above; the morphological differences
make sense in adaptive terms; wherever the taxa
meet (at the junction of bank-edge and bank-
interior coasts ) , they hybridize in narrow zones
that exhibit characteristic morphometric and ge-
netic patterns. We intend to use this combined
morphometric and genetic study as a model for
our biological revision of the entire genus.

We measured 20 characters in samples of 20 shells
(when a\ailable) in each of 52 samples spanning the
range of phenotypes and their geographic distri-
bution in Little Bahama Bank cerions. Three

^ Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Mass. 02138.

" Department of Biological Sciences, Purdue Uni-
versity, West Lafayette, Indiana 47907.

* Published and supported in part by a grant
from the Wetmore Coles Fund.

factor axes encompass nearly all infonnation (96.3
per cent) in the matrix of mean sample vectors; two
axes account for 88 per cent. Ribby and mottled
samples from Abaco sort unambiguously on the
first two axes; the third axis distinguishes mottled
samples from Grand Bahama by their characteristic
covariance ( high narrow shells with small and
niunerous whorls ) . All samples, defined as hybrids
by their geographic position in zones of interaction
( not by their morphology ) , have intennediate pro-
jections on the first two axes and plot in the inter-
mediate phenetic field between them on a triangu-
lar diagram; samples of the hybrid zone at Rocky
Point plot in perfect geographical order. Patterns
within morphotypes are equally smooth and simple.
Trend surface analysis displays the even variation
in size ( a multivariate compound of all measures )
for mottled samples throughout Grand Bahama, the
previous basis for three discrete "species"; minor,
but consistent, differences characterize slightly iso-
lated regions on Abaco — samples at Treasure Cay,
for example. Samples from areas of interaction are
intermediate in phenotype between ribby and
mottled "parental" populations. At Rocky Point,
the very narrow ( less than 1 km. ) hybrid zone dis-
plays continuous transition in mean phenotype with
no increase in variability within samples.

A study of allozxine \ariation at 28 loci (6 vari-
able and scorable ) for the same samples yields
very little concordance between biochemical data
and i^atterns of variation in shell phenot}/pes.
Cerion, though facultati\ ely hermaphroditic, are
outcrossing and moderately variable for structural
genes sur\eyed (polymorphic loci per population,
20-36 percent; average heterozygosity per indi-
vidual, 5-12 percent ) . All samples are markedly
similar. Nei's I for 820 painvise comparisons
ranges only from 0.9451 to 0.9999 (average of
0.9849); no "marker" gene characterizes any re-
gion or moipliot\pe — though characteristic fre-
quencies of \ ariable alleles clearly separate Grand
Bahamian from Abaconian populations in a statisti-

Bull. Mus. Comp. Zool., 148(8): 371-415, September, 1978 371

372 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

cal manner. The genetic analysis of h>biid samples
affirms our decision to treat the Little Bahama
Bank cerions as two semispecies, rather than as
geographic \ariation in a single entity. Although
we find no increased \ ariation in shell phenotx'pes,
hybrid Cerion are significantly more \'ariable geni-
cally (both within and among samples) than ani-
mals collected elsewhere. They are also poly-
moiphic for alleles not found in either adjacent
"parental" population.

I. INTRODUCTION

The ciiricnt taxonomy of the Little Ba-
liania Bank cerions is a microcosm of the
problems tliat beset this entire fascinating
genus, with its 600-odd named taxa (Clench,
1957; Mayr and Bosen, 1956). Little Ba-
hama Bank was spared from visits by the
most exuberant conchological splitters, but
even its conservative monographers (Clench,
1938, for example) followed the hallowed
tradition of naming every distinctive allo-
patric morphology. Seven species are now
recognized for the islands of Little Bahama
Bank.

A taxonomic scheme is not merely a
neutral description of diversity; it is, as
Mayr (1976) has emphasized, a theory of
resemblances. And, like all theory, it chan-
nels thought along prescribed lines. In
Cerion, the geographic mapping of de-
scribed taxa yields a "crazy-quilt" (Mayr
and Bosen, 1956 ) of disordered distribution.
Published reports and museum specimens
show this pattern for the seven taxa of the
Little Bahama Bank (Fig. 1). All leading
students of Cerion have invoked the vaga-
ries of hurricane transport as an explana-
tion for this incoherence (Maynard, 1919;
Bartsch, 1920, p. 53; Clench, 1957; Mayr and
Bosen, 1956). Yet if the taxonomy is incor-
rect— if these "species" are only local demes
of i^ersistent and widespread biological
species — then this biogeographic postulate
falls.

The few scientists who approached Ce-
rion with the integrative goals of modern
evolutionary biology have realized that
something in the state of its systematics must
be very rotten (Clench, 1957; Mayr, 1963;

see also Plate, 1906 and 1907 for similar in-
sights from a non-Darwinian evolutionary
perspective). As a primary though generally
unrecorded fact, no unambiguous case of
sympatry has ever been reported among
Cerions 600-odd taxa. The two most prob-
able cases are both in doubt. Mayr (1963,
p. 398) reported two species from one of
his Cuban localities, but his specimens (S.
J. Gould, personal observations) include a
few clear intermediates. Bartsch (1920) re-
ported no hybridization between two "spe-
cies" from Andros Island transplanted to the
same locality in the Florida Keys. But he
later came to question his own observation
(Bartsch, 1931, p. 373). In our own field
work, extending over five years and as
many major islands, moiphotypes ( "species"
of previous authors) hybridize freely at
their zones of contact, no matter how dis-
tinct their morphologies — and some of the
zones on Long Island mark the smooth mix-
ture of the most distinctly different mor-
phologies within the genus (e.g., smooth,
squat "C. malonei," with a long, triangular
member of the peculiar subgenus C. ( Um-
])oni.s); see Gould, Woodruff, and Martin,
1974, Fig. 1, upper row, specimens 3 and 4).
Moreover, we have detected very little ge-
netic difference among animals of diver-
gent shell morphology (Gould, et al., 1974,
Woodruff, 1975a,b). Cerion seems to pos-
sess a remarkable capacity (among animals)
for developing localized, highly distinct
morphologies without attendant reproduc-
tive isolation from other demes.

We wish to emphasize that our quest for
a revised taxonomy is not motivated by any
abstract desire for tidiness or simplification.
Bather, a more adequate nomenclature both
arises from and potentially leads to a better
evolutionary understanding of Cerions un-
usual biology. A well-revised taxonomy is
both a precondition and a promise.

We began our work in 1972 in the basic
tradition of evolutionarv natural historv.
We wished, first of all, to study selected
islands in detail and, to map the distribution
of morphological variation, hoping to find

Natural History Cerion Mil: A Revision • Gould and Woodruff 373

ABACOENSE
B ENDALLI
CHR YSALO IDES
LUCA YANORUM
MAYNAR D I
OWEN I

Figure 1. Distribution of Cerion on Little Bahama Bank as recognized taxonomically at the time this study was
initiated. M marl<s Duck Cay, suspected by Clench as being the type locality of C. milleri. Pilsbry and Vanatta
did not specify a locality for C. abacoense (beyond simply "Abaco" itself); we have placed it on the only part
of Abaco where shells of its morphology occur.

some correlation with local en\ir()nnient.
We also wanted to record everything we
could observe about the \'irtually unknown
basic biology of these snails (feeding habits,
predators, etc. ) . Beyond this, we decided to
apply a dual strategy of genetic and mor-
phometric study of the same animals (as
fruitfully applied, for example, by Soule,
1976 and Johnston, 1975). Consequently,
we collected large samples at many locali-
ties— either by gathering all the adult speci-
mens we could find in about 30 minutes, or,
in areas of high abundance, by recovering
100-200 specimens within an area of less
than 100 m-. Our genetic methods are de-
scribed in Woodruff, 1975b; our morpho-
metric approaches in Gould et al., 1974.

In our first report (Gould, Woodruff,
and Martin, 1974), we showed that a local
set of populations on Abaco Island, clearly
distinct enough morphologically to win spe-
cific designation by all previous criteria,

could only be ranked as a well-marked
geographic variant within the only taxon
inhabiting its general area. We now extend
this approach to consider the entire Cerion
fauna of Little Bahama Bank

Fig. 2;

II. GEOGRAPHIC DISTRIBUTION AND
TAXONOMIC SIMPLIFICATION

Of the two major platforms that include
most of the Bahama Islands, Little Bahama
Bank is the smaller and more northerly. It
includes ( Fig. 3 ) the two major land masses
of Abaco and associated islands on the
northeast and eastern part of the bank, and
Grand Bahama on the southwest and south.
In contrast with Great Bahama Bank (6
major islands, hundreds of minor ones and
about 250 recorded species of Cerion), it
represents a tractable area for the study of
Cerion over a broad and distinct portion of
its range.

374 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Figure 2. Representative specimens displaying the ran
ern Bahamas. Top: ribby morphotype. Bottom: mottle
row, left to right: C, chrysaloides. Grand Bahama; C. lu
ern end of Abaco, locality 250; C. abacoense, southeast
ern end of New Providence Island; C. salinaria, Salt Cay
left to right: C. bendalli, Grand Bahama, locality 200
of Great Abaco, locality 217; shell that could be assign
Providence Island, locality 267; shell that could be ass
dence, locality 275; holotype of C. degeneri from New P

ge of variation within the two morphotypes of the North-
d morphotype. Conventional taxonomy as follows; Top
cayanorum. Mores Island (holotype); C. maynardi, south-
ern shore, Abaco. locality 254; C. glans coryi from west-
north of New Providence (holotype). Bottom row, from
C. bendalli. Abaco locality 228; C. bendalli, western tip
ed to any one of 10-15 species, Culbert's Point, New
igned to any one of 10-15 species, central New Provi-
rovidence.

Natural History Cerion VIII: A Revision • Gould and Woodruff 375

EIGHT MILE ROCK

(243) (245) (244)y 4 iL^tJE CAY

ROCKr POINT

C. ABACOENSE

C. BENDALLI

INTERMEDIATES

HOLE IN THE WALL
250]

[520
B21

Figure 3. Distribution of Cerion on Little Bahama Bank revised in accordance with this study. Numbers refer
to the authors' field localities and samples (see appendix). The edge of the bank is indicated. For details of the
area of interaction on Great Abaco, see Figure 5.

Both previous monographers of Little
Bahama Bank Cerion recognized that its
several species could be allocated to two
groups within the subgenus C. (Strophiops)
(Pilsbry, 1902; Clench, 1938). Beyond this
basic statement, the literature contains
nothing of an explanatory or integrative
nature. We have only a list of localities and
taxa.

The two groups are distinct in moq^hol-
ogy. Shells of the "ribby" morphotype are
white or weakly mottled, relatively wide,
and cylindrical with a fairly sharp break be-
tween a triangular apex and parallel-sided
later whorls, strongly recurved aperture
with thick lip, and a complete covering of
strong, often widely spaced ribs (Fig. 2) —
in short, a lightly colored, heavy and ribby
shell. Shells of the "mottled" morphotype
are strongly colored with irregular, brown-
ish mottling, generally narrow with a more

rounded apex passing smoothly to more
barrel-shaped later whorls, apertures either
thickly or thinly lipped depending upon the
habitat (though never so thickly lipped as
the ribby morphotype ) , with a shell surface
either smooth or covered with fine ribs
(Fig. 2) — in short, a mottled, light and rela-
tively smooth shell.

We use the archaic term "morphotype" to
describe these basic features because we
find the same contrast — and the same corre-
lation with geographic position and habitat
— on island after island in the northern
Bahamas. W^e have no reason to assert
homology and transport among islands,
though this has been the unstated assump-
tion of all previous work. It is just as likely,
we believe, that these basic morphologies
are developed in situ, again and again, as
adaptive responses to recurring habitats.
On each island, the ribbv and mottled

376 Bulletin Muscutn of Comparative Zoology, Vol. 148, No. 8

morphotypes interbreed in zones of contact;
yet all the zones display features (varying
from island to island) suggesting that at
least a minor amount of genetic differentia-
tion has occurred. They are imperfectly
separated forms, perhaps best designated
as semispecies, if conventional categories
must be applied. (The biological species
concept breaks down for an animal like
Cerion with such amazing morphological
diversity accompanied by, at best, imper-
fect reproductive isolation. We can scarcely
recognize but a single species for a pat-
tern of discrete and coherent morphological
xariation unexcelled among genera of land
snails. Yet we cannot identify taxa by
lack of interbreeding in sympatry.) We
envisage a basic genetic system, common to
all Cerion and including the potential to
develop any one of a set of basic morpho-
types. The morphotypes are alternative
pathways of development that can be
evoked from a common genotype by mecha-
nisms of regulation utterly unknown to us.
Once evoked, however, these morphotypes
can become relatively stable within local
areas. The basic features of any morpho-
type do not form a labile ecophenotype,
easily altered by rearing in different condi-
tions (Bartsch's transplants of Bahamian,
Cuban, Puerto Rican, and Curasao cerions
all bred true to type for at least two genera-
tions on the Florida Keys and Dry Tortugas
— Bartsch, 1920).

We believe that a modern taxonomy of
the Little Bahama Bank cerions can do no
more than recognize the two morphotypes
as imperfectly separated semispecies. We
base this conclusion on three sets of obser-
vations: consistent geographic distribution
of the morphotypes, adaptive correlations
with habitat, and patterns of interaction in
zones of contact.

1. Geographic distribution. Among the
myths that surround Cerion, none has been
more persistent than the claim that it is a
halophilic species restricted to coastal areas.
All previously reported records for both
ribby and mottled morphotypes are from

localities within about 100 m of the sea. Yet
we have found that the mottled morphotype
ranges right across these low islands, pene-
trating the middle of the Grand Bahamian
pine forest and the middle of the once for-
ested area of Abaco. At locality 204 (see
Fig. 3), 10 km from the nearest coast we
found mottled Cerion at very low densities
(< 0.1/m-) in the shrubs and grass on the
forest floor. Beneath one slab of aeolianite,
however, we discovered an aggregation of
more than 50 adults. Only in open, dis-
turbed areas in the forest (Iocs. 205, 207)
did we find Cerion in abundance (approx.
1/m-), and, even then, not in numbers
typical of coastal populations where densi-
ties greater than 10/m- are common. These
sparse and patchy interior populations un-
doubtedly escaped the notice of early col-
lectors, whose activities were usually re-
stricted to a few minutes walk from the
point where they beached their dinghies.
W. J. Clench (1938), the most careful col-
lector of Cerion found some interior speci-
mens, but did not appreciate the generality
of their occurrence.

In contrast to our discovery that the mot-
tled morphotype ranges far from present
day coasts, the ribby morphotype is re-
stricted to within 200 m of the coast. Fur-
thermore, and most importantly, it is re-
stricted to coastal areas adjacent to the edge
of the island bank (Fig. 3, for example). In
contrast, the mottled morphotype occurs
along coasts that do not border the island
bank. If we designate the ribby morpho-
type as having a "bank edge" distribution,
then the mottled morphotypes are found in
"bank interior" situations. The mottled
shells may represent an inland or bank in-
terior morphotype evolved for geographic
or ecological conditions prevailing during
Pleistocene hypothermal periods when the
sea level was much lower than it is today. If
this hypothesis is correct, then the mottled
morphotype has been living in coastal situ-
ations (along the northern coast of Grand
Bahama and the western coast of Abaco)
for less than 6,000 years. In contrast to the

Natural History Cerion VIII: A Revision • Gould and Woodruff 377

traditions of Cerion study, and for reasons
presented herein, we believe that current
I distributions may be highly persistent. The
" preference for fluid, haphazard distribu-
. tions proposed by earlier workers (illus-
I trated in Fig. 1) arises from a taxonomy
that we will show to be fundamentally in-
correct.

Distribution patterns based on the revi-
sions in this paper are shown in Figure 3.
The generally coherent pattern of bank
edge \'s. bank interior distribution found in
these two taxa is one of our most important
findings: it permits us to predict the dis-
tribution of analogcms morphotypes on the
\'arious islands of the Great Bahama Bank.
On Andros, New Providence, Great Exuma,
and Long Island, we have found that the
mottled morphotype invariably lives on
bank interior coasts and inland areas, while
the ribby morphotype is restricted to bank-
edge coasts. We expect eventually to show
that more than 200 "species" of Bahamian
Cerion only represent the distribution of
these two moiphotypes and their interac-
tion.

The consistent differences in distribution
provide, in themselves, a strong argument
for regarding the two morphotypes as partly
distinct, biological taxa. They live on dif-
ferent kinds of coasts and react differently
to inland conditions. Were it not for their
patterns of interaction (see below), we
might regard this strong correlation of form
with habitat and geography as an aspect
of normal geographic variation within a
single taxon (perhaps purely phenotypic),
rather than as the adaptations of imper-
fectly separated entities.
j 2, Adaptation of form to habitat. For all
the effort devoted to taxonomy ( more than
2,000 printed pages), no previous workers
have directly studied the adaptive nature of
form in Cerion. Nonetheless, the persistent
correlation of form and habitat suggests
that the morphotypes have been selected for
survival xalue. Accordingly, we have initi-
ated a series of experiments designed to
establish some of the physical correlates of

the various morphologies. Looking first for
the possible adaptive significance of shell
pigmentation, we contrasted the white shells
of the ribby morphotype with those of the
mottled moiphotype. John Quensen, work-
ing in Woodruff's laboratory, found that in
direct sunlight the interior of a mottled shell
averages 1 C warmer than the interior of an
unpigmented shell. It may well be that the
ribby shells, characteristic of exposed bank-
edge situations, are protected from over-
heating by the lack of shell pigmentation.
Such an ecological correlation between
shell color and body temperature has been
found in other land snails (Rensch, 1932;
Schmidt-Nielson et al., 1971; Yom-Tov,
1971; Heath, 1975). It is also possible that
shell pigmentation plays a role in predator
avoidance. The mottled shells are initially
hard to find, as they hang from bush stems
and on blades of grass in the dabbled sun-
light and shadow ( a clear case of disruptive
coloration to our eyes) (Fig. 4). In con-
trast, the white shells of the ribby morpho-
type are fairly conspicuous on the stems and
leaves of bushes and other plants. Only
when they descend to the ground in rocky
areas is their coloration at all cryptic. In a
second investigation, Quensen has examined
Vermeij's (1975) suggestion that sculptur-
ing (ribbing) is a defensive adaptation in
snails since it confines the predator's crush-
ing force to the thickest part of the shell.
Quensen's preliminary results indicate that,
in Cerion, overall shell size is more impor-
tant than ribbing in determining the
strength of the shell. Approximately SO per-
cent of a shell's ability to resist fracture is
attributable to shell weight and shell height;
interrib shell thickness is more significant
than shell thickness at a rib or ribbing den-
sitv. This does not mean that ribs are
unimportant in Cerions defense, but only
that they do not protect the animal from
compression applied generally along the
sides of the shell. While the identity of
Cerions key predators remains unknown,
\\'Oodruff's detailed population studies on
Abaco and elsewhere implicate land crabs,

378 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Figure 4. Cryptx nature of mottled coloration. When sunlight is filtering through bushes, the mottled shells
are very hard to see (at least for us). Photo taken by J. IVIartin on northeast coast of Great Abaco.

rats, and possibly a bird. The results of
these studies will be reported elsewhere
(Woodruff and Quensen, in prep.).

3. Patterns of interaction. Populations of
the ribby niorphotype once inhabited the
bank edge at Eight Mile Rock on the south-
west Coast of Grand 13ahama ( Plate, 1907 ) .
In 1936, Clench and Greenway searched ex-
tensively for this form in the area where
Millspaugh originally collected it. After two
weeks they found only a single dead shell on
the eastern side of Hawksbill Creek. Clench
(1938) concluded that the hurricane of
1935 may have destroyed this colony, as it
did a great deal of damage along the entire
south coast of the island. In 1963 and 1964,
small samples of ribby shells were again
found at Freeport and Smith's Point ( speci-
mens in the Museum of Comparative Zool-
ogy). In September 1972, we spent several
days searching the south coast of the island.

from Freeport to West End; no Cerion were
found. This is the only case we know in
which a niorphotype has apparently become
extinct on an entire island.

On Abaco, however, we need only a map
of bank edges to predict exactly where the
contacts between ribby and mottled mor-
photypes should occur. The village of
Sandy Point ( Fig. 5 ) marks the coastal
transition from bank edge to bank interior;
here we collected a sample of intermediate
morphology. The ribby niorphotype inhab-
its the coast south of Sandy Point, around
the southern tip of the island, up to the
narrow area known as The Crossing (Fig.
5). Here, the second contact occurs as the
interior, mottled populations are squeezed
into close contact with ribby animals on the
eastern shore. We observed no interactions;
a narrow hill, running parallel to the coast,
seems to separate the morphotypes com-

Natural History Cerion VIII: A Revision • Gould and Woodruff 379

pletely. Yet there must be some "leakage"
across the hill, for morphometric analysis
(see below) demonstrates the intermediate
nature of apparently mottled shells at this
locality. According to the map, we must
predict an interaction around Cherokee
Harbor (Fig. 5), for here the coast again
switches from bank edge to bank interior.
Here, indeed, is the third and by far the
most interesting interaction. Ribby popu-
lations, extending from the south, encounter
mottled populations from the north in an
apparently smooth, but local hybrid zone.
We shall analyze this zone in detail in the
following sections on morphometries and
genetics (see also Woodruff and Gould, in
press ) since it holds the key to our interpre-
tation of these two taxa.

The current taxonomy of Abaco cerions
recognizes seven taxa within these two mor-
photypes. Ribby populations have been al-
located to four species :

1. Cerion abacoense Pilsbry and Vanatta,
1895, p. 209. The type specimen (Acad.
Nat. Sci. Phila. No. 25337) and all associ-
ated museum material (M.C.Z., Harvard
University, and Acad. Nat. Sci. Phila.)
clearly indicate that this name applies to
ribby populations from The Crossing, north
to the hybrid zone. These shells are some-
what smaller, lighter with more though
weaker ribs than southern samples. Pilsbry
and Vanatta list their locality simply as
"Abaco Island."

2. Cerion maynardi Pilsbry and Vanatta,
1895, p. 210. Again, listed only as "Abaco
Island," but we have found this morphology
only near Hole-in-the-Wall Light near South
Point (southern tip of the island) where
ribby shells are larger with very strong and
sparse ribs.

3. Cerion chnjialoides Plate, 1907, p. 597.
The extinct, bank-edge population collected
by Millspaugh at Eight Mile Rock on the
southwestern coast of Grand Bahama.
Shells are considerably smaller than those
of other ribby populations, but differ from
them in no other evident way.

4. Cerion hicaijanorum Clench, 1938. A

longer and more slender shell with more
numerous and finer ribs. From Mores
Island (Fig. 3).

These populations are distinct in mor-
phology one from the other. Indeed, all
students of Cerion (including ourselves)
agree that virtually every local population
in this peculiar genus has its own recogniz-
able form. ( Disagreement centers only on
appropriate taxonomic definition.) In this
case, we cannot possibly justify any sepa-
ration into species. We can barely distin-
guish the far more different ribby and
smooth populations on the basis of their
patterns of interaction. It is not likely that
any reproductive barriers exist among lo-
cal populations of the same moiphotype.
(Gould and Paull, 1977, have lumped
within-morphotype variation for all cerions
from Hispaniola to the Virgin Islands into
a single species.) ^^'e therefore reject C.
maynardi, C. chrysaloides, and C. lucay-
anoruni as synonyms of the first-named
form, Cerion abacoense. The ribby morpho-
type of Little Bahama Bank should bear
this name, at least until we can determine
whether it is homologous with populations
of the ribby morphotype on islands of
Great Bahama Bank.

At least two, and possibly three, names
are available for populations of the mot-
tled morphotype.* The rejected names for
the ribby morphotype apply to geographi-
cally distinct subpopulations meriting sub-
specific rank, if we were inclined — as we
decisively are not, lest Cerion maintain its
burden of hundreds of names — to use this
category. The "species" of the mottled mor-
photype, on the other hand, have no
geographic definition; they are names

* Things could have been worse. Specimen
labels in the Department of Mollusks, United States
National Museum, include two additional names,
apparently never published by Bartsch. These
anagrams of the island — C. mahaba (U.S.N.M.
No. 179439) and C. hamaha (U.S.N.M. No.
369715) — both apply to dwarfed forms of C.
hendalli inhabiting the northern coast of Grand
Bahama Island.

380 BuUetin Museum of Comparative Zoology, Vol. 148, No. 8

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0 Ml

Figure 5a. Area of interaction between C. bendalli and C. abacoense on southern Great Abaco. Modal mor-
photype at each locality is indicated: C. bendalli, open circle; C. abacoense, closed circle; intermediates, half-
closed circle.

for minor, recurrent differences in form
througliout tlie range of mottled denies:

1. Cerion bendalli Pilsbry and Vanatta,
1896, p. 332. In an uncharacteristic act of
lumping ( overlumping, in our judgment!)
Pi]sl)ry and \'anatta originally defined C.
bendalli as a subspecies of the ribby C.
abaeoense — though they wrote (1896, p.
333): "This form at first sight looks ex-
tremely different from C abacoense, and as
we have seen no intermediate examples, it
may well prove to be a distinct species."
In 1902, Pilsbry returned to his former con-
sistency and elevated C. be}idalli to specific

rank. Pilsbry and Vanatta applied this
name to samples of the mottled morphotype
with fine ribs.

2. Cerion oweni Dall, 1905, p. 443. A
name for smooth or very finely ribbed
samples of the mottled morphotype; no
other characters distinguish it from C. ben-
dalli. Dall ( 1905) recognized three subspe-
cies within C. oueni (C. oweni incisiini, Co.
verniicuhini, and Co. veticulatum) , but
these have already been rejected by Clench
(1938, p. 328).

3. Cerion milleri (Pfeiffer), 1867, p. 129.
Pfeiffer applied this name to a small sample

Natubal History Cerion VIII: A Revision • Gould and Woodruff 381

Figure 5b. Area of interaction between C. bendalli
and C. abacoense on southern Great Abaco. Modal
morphotype at each locality is indicated: C. bendalli.
open circle; C. abacoense. closed circle; intermedi-
ates, half-closed circle.

of mottled shells from "Duck Cay, Exuma
Group, l^ahama Islands." But Clench (1933,
p. 50) noted that the Exumas contain no
Duck Cay, while an appropriate islet of this
name sits in Cherokee Harbor, Abaco. He
therefore supposed that C. milleri might be
an Abaconian species. If Clench is correct,
then C. miUeri, as the oldest available
name, should designate the mottled cerions
of Little Bahama Bank. Yet we prefer to
leave it in limbo, for we do not know how to
verify Pfeiffer's locality; mottled shells are
much of a muchness throughout the Ba-
hamas, and occur throughout the Exumas.
We therefore reject C. oiceni Dall (with
its three subspecies) and C. milleri (Pfeif-
fer) and designate the mottled morphotype
on Little Bahama Bank as Cerion ])endaUi
Pilsbry and \'anatta, 1896.

III. MORPHOMETRICS OF CERION ON
LITTLE BAHAMA BANK

A) Introduction

We were originally attracted to Cerion
because it is such an ideal animal for mor-
phometric study. Like most mollusks, it
preserves a complete record of its ontogeny
in an accretionary shell. Its particular ad-
\'antages arise from two properties of
growth : 1 ) The transition between em-
bryonic shell and later accretionary growth
is precisely marked by a discontinuity in
ribbing and rate of expansion; we therefore
obtain an unambiguous, biological criterion
for numbering whorls; we take this dis-
continuitv as the beginning of the O'th
whorl. This numbering permits us to define
morphometric properties at a variety of
standardized stages throughout growth. 2)
As it reaches maturity, Cerion changes its
direction of coiling and, finally, secretes a
terminal adult aperture with an expanded
and reflexed lip. We can therefore measure
the traits of its definitive adult size. (Most
mollusks have no stage of terminal growth;
we can define neither the mean nor variance
of adult characters because we cannot sort
ontogenetic from static adult variation.) In
Cerion, we can compare adult characters
with corresponding traits at any stage of
growth; in most mollusks, we can define
neither set of measures unambiguously.

We have chosen a suite of variables that
should measure all of the traits (except
color) commonly used to erect taxa within
Cerion. Our set also defines the major as-
pects of growth and co\'ariation: size and
shape of the embryonic shell, patterns of
ribbing, size and shape of juvenile and pre-
adult whorls, number of whorls, measures
of final size, and characters of the adult
umbilicus and aperture. Although our mea-
sures contain some inevitable redundancy,
our previous studies clearly demonstrate at
least five independent patterns of covaria-
tion among them (Gould et al., 1974; Gould
and Paull, 1977).

Our measures follow the definition and

382 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

protocol of Gould et al. ( 1974, pp. 522-524)
with the exception of 6 and the addition of
20 (used only as the numerator of ratio
measure 18 in Gould et al., 1974; we have
since determined that it includes interesting,
independent information of its own):

1. width of the protoconch

2. width at the end of the fourth whorl

3. total number of whorls (with the
termination of the protoconch taken
as the O'th whorl)

4. number of ribs on the fourth whorl

5. number of ribs on the sixth whorl

6. number of ribs in 50 micrometer
units at the termination of the first
whorl

7. length of the adult shell, apex to
lower apertvual tip

8. maximum width of the adult shell

9. height of the protoconch

10. total height of the shell at the end of
the fointh whorl

11. height from the end of whorl 4 to
the end of whorl 6

12. width of the umbilicus

13. width of the apertural lip at its
widest point (measured parallel to
the plane of the aperture

14. thickness of the apertural lip at its
thickest point (measured perpendic-
ular to the plane of the aperture)

15. height of the aperture

16. width of the aperture

17. protrusion of the aperture

18. tilt of the aperture

19. weight of the shell

20. distance from aperture to preceding
suture: line EG of Gould et al.,
1974, fig. 5, p. 523.

B) The Basic Pattern

We chose 52 samples, representing all
taxa and habitats, and measured 20 shells
from each sample when available — 14 sam-
ples contain fewer shells, but only 5 of
these have fewer than 15 specimens.
Localities are listed in the appendix. Forty-
eight samples are from our own field collec-
tions, 4 from the collection of the Depart-

ment of Mollusks, Museum of Comparative
Zoology, Harvard University [3 of the ex-
tinct ribby morphotype ("C. chrysaloides")
from Grand Bahama, 1 of "C. lucaijanorum"
from Mores Island]. We are more than
conventionally grateful to John Hevelin for
spending half a year compiling one of the
most scrupulously accurate data sets ever
assembled in molluscan biometrics.

Many strategies are available for reducing
a data set of 20 measurements on nearly
1000 specimens from 52 samples. We de-
cided to treat each sample as a potentially
random extract from a single statistical uni-
verse, rather than as a definite entity to be
separated, if possible, from other groups.
This decision — a methodological correlate
of our belief that Cerion is a single entity
with local inhomogeneities led to a factor-
analytic model. We computed the mean
vector for each sample (Table 1) and per-
formed a Q-mode factor analysis of the 52
items using program CABFAC (Klovan
and Imbrie, 1971 ) . We included the follow-
ing data transformations :

1. percent-range method of equalizing
weights. The highest value of each variable
receives a value of 100, the lowest becomes
0; others are scaled as a percentage of this
range. This is not always (or even often)
a desirable method for achieving equality of
weights. Suppose, for example, that a trait
varies narrowly and randomly among speci-
mens. We would not want such variation
to count as much as the wider range of
another measure clearly adapted to vari-
ation in habitat. But, in this case, our
values are well-determined means of sam-
ples, not the random error of individual
specimens. A stable narrow range may be
just as important as a wider one.

2. normalization of vectors. Each vector
is rescaled to unit length before the extrac-
tion of eigenvalues. This transformation re-
moves the explicit influence of variation in
average shell size among samples. ( How-
ever, the allometric correlates of size may
still be expressed as shape.) We preferred
to eliminate this explicit variation in size

Natural History Cerion VIII: A Revision • Gould and Woodruff 383

Figure 6. Position of mean vectors for all samples of Little Bahama Bank cerions. This is a triaxial plot of nor-
malized factor loadings for a 3-axis, varimax solution in the Q-mode. These three axes explain 96.3 per cent of
all information; the first two axes explain 88 per cent. Ribby and mottled morphotypes are well separated by
the first two axes. Mottled samples from Grand Bahama Island have higher projections on the third axis. Closed
circles are mottled samples from Abaco; open circles are mottled samples from Grand Bahama; crosses are
ribby samples; squares represent samples defined by geography and ecology (not morphology) as inhabitants of
zones of interaction between ribby and mottled populations (note their intermediate position in morphology as
well); the star represents the single fossil sample from Abaco. The line connects samples of the hybrid zone at
Rocky Point in geographical order, pc is the Pongo Carpet sample (mottled, partly convergent upon ribby); f is
the fossil sample; cry are "C. chrysaloides" (the name applied to ribby samples on Grand Bahama); luc is "C.
lucayanorum" (ribby sample from Mores Island). Other numbers refer to localities discussed in text.

because it can control so much covariance in
a matrix (large shells have high values of
almost all variables), and because all mor-
photypes and areas contain both large and
small-shelled samples. We eliminate this
pervasive control of size in order to see
smaller but more stable influences more
clearly. However, we also performed an
analysis without normalization and obtained
nearly identical results (see below).

Three axes encompass 96.3 per cent of
the information in 52 samples; no subse-
quent axis reaches one per cent. We per-
formed a varimax rotation and computed
the factor loadings of all samples upon the
three axes ( in Q-mode analysis, samples are

loadings ) . CABFAC normalizes the triaxial
loadings to permit a plot as a triangular
graph.

Figures 6-7 display a remarkable result.
All the variation in Little Bahama Bank
cerions, the basis of 7 species and a host of
subsidiary distinctions, reduces to a matrix
not far from rank 2! (Two varimax axes ex-
plain 88 per cent of all infoiTnation.) And the
foci of these axes are our two old friends —
the ribby and mottled moiphotypes in their
"pure" form. All intermediate samples from
zones of contact — and only these samples —
plot in between. Moreover, the minor tliird
axis has its own coherence, for all samples
that load strongly upon it (with one excep-

384 Bulletin Musciu)} of Comparative Zoology, Vol. 148, No. 8

Table 1. Matrix of means (in mm, g, or couxts) for all samples treated biometrically in this
WORK. (Converted from original data in micrometer units — variables 1, 9, 13, 14 multiply by
18.0; VARIABLES 2, 10, 11, 12, 15, 16, 17, 20 multiply by 8.0 for micrometer units. All biometrical
WORK done in micrometer units. Data in this form available from authors. )

Proto-

Sami^le

conch

4th whorl

total

conch

Number Location width

width

whorls

4th ribs

6th ribs

Lst ribs

height

width

height

92367 Grand Bahama: 2.74

7.79

7..50

21.33

19.67

6.83

22.77

9.57

1.26

247236 ribby

2.84

8.16

7.41

22.85

20.80

7.08

23.90

10.11

1.38

250620

2.78

8.23

7.33

23.47

21.67

7.63

23.21

9.96

1.39

212 Grand Bahama: 2.93

S.20

8.30

79.83

57.60

17.00

23.99

9.05

1.19

211 mottled

2.94

8.50

8.71

48.00

44.00

11.45

26.96

10.44

l.,30

208

3.03

8.81

8.29

60.60

52.20

12.53

25.13

9.81

1.26

204A

3.29

9.65

9.20

53.83

47.78

12.58

,32.09

11.86

1.39

204B

3.15

9.41

8.63

48.75

44.05

12.05

27.86

11.28

1.27

207

3.09

8.22

9.28

86.00

77.95

16.13

28.91

10.79

1.35

209

2.79

7.52

8.99

100.00

81.00

13.58

23.05

9.17

1.27

205

3.11

9.25

8.80

73.10

68.80

12.42

29.86

11.53

1.25

202

3.19

9.47

8.51

71. .30

65.65

13.65

29.74

11.61

l.,39

199

2.71

7.27

7.20

94.88

73.31

16.83

20.07

8.08

l.,32

200

3.15

8.37

8.71

99.25

78.50

18.10

27.11

9.93

l.,32

201

3.03

8..33

8.93

93.90

83.35

15.60

27.68

10.18

1.27

213 Little Abaco 3.22

9.36

8.70

85..55

75.55

15.78

31.27

11.06

1.32

214

3.13

9.14

8.38

100.85

86.65

17.93

28.05

10.58

1.22

216

3.04

8.81

8.17

93.70

73.05

17.85

26.15

10.08

1.29

217 Great Abaco 3.17

9.18

8.21

95.56

71.28

16.09

28.83

11.00

1..35

218

3.00

8.98

9.04

80.25

60.40

14.85

32.61

11. ,37

1.28

316

2.85

8.03

7.93

58.45

41.15

13.85

24.68

9.63

1.34

228

3.02

8.31

7.89

81.45

63.10

14.43

24.27

9.61

1.50

229

3.05

8.39

8.02

79.21

67..33

15.97

25.31

9.87

1.47

230

3.00

8.08

8.43

95.95

85.21

18.23

26.08

9.82

1.41

231

3.05

8.74

8.85

78.80

61.10

14.82

,30.49

11.47

1.48

233

3.42

9.06

9.25

96.00

76.00

16.50

31.10

11.20

1.54

240

3.10

8.89

8.71

80.65

62.65

15.72

29., 56

10.83

1.27

247

3.18

8.81

7.88

103.20

75.35

17..30

25.26

10.15

1.26

246

3.02

8.38

8.42

47.68

40.00

10..58

25.92

9.93

1.55

243

3.29

8.97

8..53

100.00

84.06

17.91

27.08

10.61

1.31

245

3.38

9.66

8.23

95.70

79.35

17.05

28.14

11.00

1.37

244

3.25

9.17

7.79

97.80

74.85

14.90

25.23

10.24

1.24

261

3.21

9.73

8.55

94.71

78.57

15.36

30.26

10.99

1.33

260

3.13

9.40

7.84

58.85

49.45

12.63

26.73

11.13

1.31

308

3.08

9.23

7.68

53.50

45.65

12.90

25.76

10.96

1.31

307

3.02

9.38

7.84

46.65

41.10

11.40

26.97

11.39

l.,35

306

3.17

9.95

7.93

44.10

38.90

10.95

28.68

12.18

1.35

310

3.39

10.53

8.13

95.82

74.90

15.78

27.60

11.82

1.39

309

3.38

9.78

8.12

98.33

89.56

16.70

28.88

11.38

1.37

305

3.12

9.95

7.76

,39.80

34.65

10.53

28.83

12.15

1.37

304

3.11

9.82

8.08

44.40

39.90

11.43

29.52

12.30

1.33

311

3.04

9.51

7.94

44.20

.39.15

10.95

27.52

11.45

1.35

259

3.08

9.68

8.22

38.80

34.45

9.95

31.37

12.83

1.36

249

3.63

10.35

9.46

82.05

74.95

14.27

34.40

12.71

1.44

254

3.29

10.49

7.87

38.55

33.05

9.18

29.97

12.70

1.44

303

3.29

10.38

7.88

43.40

37.47

9.85

29.40

12.58

1.35

253

3.34

9.51

8.05

67.50

44.45

13.20

27.49

10.91

1.30

255

3.18

9.10

8.31

73.50

48.60

14.40

28.14

10.53

1.25

257

3.11

10.12

8.04

38.55

32.65

9.15

30.10

12.72

1.35

251

3.01

9.07

8.05

53.55

41.65

12.55

26.98

11.20

1.36

250

3.37

10.90

8.79

26.20

24.85

8.53

,34.35

13.76

1.52

LUC Mores Island 3.09

9.42

8.10

42.02

32.10

10.40

30.26

11.62

1.43

Natural History Cerion VIII: A Revision • Gould and Woodruff 385

Table 1 [continued]

4th whorl

4th-6th

nnibilical

lip

aperture

aperture-

height

width

thickness

height

width

protrusion

tilt

weight

suture

6.65

10.92

4.63

1.22

1.09

8.83

7.25

2.50

1.64

.79

4.42

6.88

11.64

4.55

1.23

1.09

9.31

7.73

2.47

1.82

.73

5.13

6.84

11.52

4.52

1.22

.88

9.05

7.42

2.10

1.82

.68

4.83

6.15

9.71

4.30

.98

.65

8.81

7.15

2.62

2.15

.49

4.51

6.08

9.87

4.64

.91

.63

9.38

8.01

2.43

1.98

.71

5.51

6.25

10.01

4.38

1.13

.73

9.12

7.41

2.54

2.05

.62

5.01

6.26

10.17

5.48

1.21

1.14

11.10

8.99

3.15

1.95

1.49

6.30

6.03

10.28

4.93

1.11

.84

10.19

8.50

2.81

2.06

.96

5.73

6.10

8.79

4.94

.95

.62

9.76

8.10

2.31

1.87

.82

5.42

5.38

7.49

4.12

.85

.44

7.86

6.63

2.13

2.21

.37

3.79

6.22

10.59

5.19

.98

.69

10.73

9.07

2.78

2.04

1.12

6.18

6.57

11.06

5.14

.99

.75

10.65

8.96

2.64

2.11

1.36

6.23

6.60

10.03

3.13

.71

.52

7.51

6.49

1.93

2.15

.41

5.05

6.68

9.53

4.49

.96

.63

9.49

7.68

2.76

2.30

.81

5.49

6.24

9.42

4.74

.99

.74

9.72

7.87

2.75

2.20

.83

5.39

6.59

11.10

5.59

1.38

1.47

11.94

9.29

3.35

2.19

1.48

6.01

6.42

10.70

5.19

1.14

.91

10.77

8.61

3.06

2.40

1.21

5.41

6.65

10.49

4.68

1.06

.85

10.18

8.21

2.86

2.28

.89

4.91

6.74

11.57

4.96

1.18

1.20

11.42

9.18

3.23

2.40

1.11

5.85

6.50

10.73

5.91

1.46

1.32

12.61

10.13

4.01

2.81

1.34

5.96

6.44

10.63

4.23

1.13

1.04

9.41

7.94

2.64

2.00

.84

5.24

6.72

10.44

4.49

1.06

.84

9.38

8.11

2.93

2.65

.62

4.82

6.69

10.63

4.81

.99

.83

9.51

8.24

2.69

2.19

.69

5.12

6.35

9.88

4.73

.90

.74

9.44

7.76

2.62

1.99

.69

5.14

6.64

10.08

6.03

.96

.83

11.16

9.08

2.74

1.86

.98

5.69

6.66

9.43

5.25

1.47

.92

11.50

9.34

2.78

2.26

1.12

5.63

6.34

10.11

5.95

1.13

1.21

11.18

9.05

3.88

2.49

1.15

5.26

6.58

10.96

3.83

1.02

1.06

9.93

8.37

2.67

2.19

1.00

5.99

7.38

10.45

4.43

.98

1.38

9.95

8.08

3.29

2.39

1.12

5.89

6.48

10.06

4.93

.96

1.11

9.93

8.31

3.14

2.31

1.13

5.28

6.83

11.58

5.13

1.06

1.17

10.58

9.11

3.15

2.15

1.15

5.88

6.89

11.18

4.43

.85

.87

9.78

8.24

2.95

2.40

.87

5.35

6.81

11.30

5.63

1.12

1.09

11.25

9.08

3.06

2.09

1.39

5.85

6.73

11.63

5.63

1.09

.84

10.73

8.71

3.00

2.03

1.06

5.44

6.77

12.08

5.16

.95

.79

10.23

8.40

2.71

2.05

.95

5.82

6.74

12.04

5.31

1.17

.89

10.59

8.97

2.82

1.95

1.13

6.20

6.84

12.42

5.98

1.20

1.07

11.28

9.23

2.98

1.85

1.58

6.27

6.47

11.73

5.56

1.14

.67

10.53

8.84

2.89

2.27

.76

5.41

6.26

10.40

5.34

1.01

.72

10.50

8.74

2.67

2.20

.86

5.58

7.04

13.28

6.18

1.15

1.04

11.76

9.31

3.18

1.83

1.71

6.15

6.76

12.16

5.81

1.32

1.25

11.95

9.72

3.22

2.37

1.70

6.53

6.62

11.73

5.60

1.21

1.01

11.21

8.95

3.11

2.20

1.08

5.84

6.84

12.73

6.71

1.54

1.61

12.91

10.29

3.51

2.41

2.13

6.24

6.27

10.30

6.19

1.15

.88

12.39

10.19

3.41

2.21

1.36

6.73

7.54

13.26

5.95

1.40

1.38

12.23

10.29

3.36

2.50

1.83

6.15

7.15

13.35

6.33

1.53

1.55

12.37

10.23

3.40

2.12

1.48

6.29

6.98

11.49

5.37

.99

1.08

10.89

9.07

3.07

2.21

1.01

5.48

6.80

10.98

5.18

.98

10.72

8.75

3.15

1.99

.98

5.59

7.20

12.87

6.23

1.61

1.15

12.53

10.43

3.24

2.14

1.80

5.89

6.80

11.41

4.79

1.15

1.43

10.66

8.93

2.95

2.31

1.34

5.82

7.20

12.11

6.62

1.93

1.84

13.14

11.23

3.73

2.12

2.07

6.41

7.36

12.48

5.89

1.55

1.60

11.71

9.77

3.49

2.19

1.54

5.99

386 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Z*-^

Figure 7. Representative shells for samples depicted in same positions on Fig. 6. At bottom line, left to right:
locality 230 (typical C. bendalli); locality 316 (convergent C. bendalli from Pongo Carpet); locality 253 (intermedi-
ate shell from The Crossing); locality 254 (C, abacoense); "C. chrysaloides" from Grand Bahama. Above: locality
204 (C. bendalli from Grand Bahama).

tion) are C. bendalli from Grand Bahama.
The third axis divides samples of C. hen-
dalli (the mottled morphotype) into its
two isolated areas.

The matrix of factor scores (Table 2)
permits us to infer the basis of separations
in Figure 6 (consult the matrix of mean
values — Table 1 — for the raw information ) .
Only three variables score highly on the
first axis. (This a.xis serves as a dimension
of reference for the mottled morphotype, C.
bcmlaUi. Samples of C. ])endaUi load
strongly upon it, and weakly upon the
second axis — see Figs. 6-7. ) Not surpris-
ingly, these three variables are the ribbing
measures 4-6. (Mottled samples always
have much weaker ribs than ribby samples,
but the ribs are always far more numerous
in mottled samples; all our ribbing measures
are counts.) No other variable so con-
sistently separates C. bendalli from C.
abacoeyise. The second axis, with it high
loadings for C. abacoense (the ribby mor-
photype), contains high scores for most

measures of final size and whorl size. ( The
scores are negative in this case. The sign
is of no particular importance, since it only
indicates the direction of the reference vec-
tor. The pattern of scores and loadings
would not change if the vector pointed 180°
in the opposite direction, thus reversing all
the signs.) To some extent, this suite of
high scores only mirrors the distinction by
ribbing made on the first axis. Since refer-
ence vectors are normalized, a small number
of ribs must lead to a greater contribution
to the vector from other measures. But the
ordering of intensity within this group of
high scores clearly distinguishes the primary
characteristics of C. abacoense. Shells of
C. abacoense do not have more whorls than
C. bendalli (note small positive score for
whorl number — primarily due to low whorl
numbers of small "C. chrysaloides"), and
they are not generally taller (modest
score for shell height). The highest scores
belong to measures of size that best distin-
guish the two taxa by higher mean values

Natural History Cerion VIII: A Revision • Gould and Woodruff 387

Table 2. Factor scores of original variables

UPON THE three FACTOR AXES USED TO DEPICT
SAMPLES IN FIGURE 6.

protoconch width

0.155

-0.102

-0.206

4th whorl width

0.070

-0.228

-0.191

total whorls

0.280

-0.007

-0.387

4th ribs

0.536

0.162

0.122

6th ribs

0.491

0.149

0.030

1st ribs

0.492

0.108

0.153

height

0.129

-0.187

-0.292

width

0.053

-0.247

-0.182

protoconch height

0.078

-0.204

0.283

10.

4th-height

0.072

-0.341

0.514

11.

4th-6th height

0.025

-0.362

0.344

12.

unil)ilical width

0.071

-0.273

-0.149

13.

lip width

-0.018

-0.236

0.041

14.

lip thickness

-0.025

-0.272

0.122

15.

apertnre height

0.062

-0.258

-0.182

16.

aperture width

0.047

-0.226

-0.175

17.

protrusion

0.087

-0.200

-0.106

18.

tilt

0.239

-0.040

0.102

19.

weight

-0.003

-0.242

-0.158

20.

aperture-suture

0.119

-0.265

-0.036

for C. ahacoense. Shells of C. abacoense are
heavier (measure 19), and wider both in
spire (S) and umbilicus (12); they have a
larger aperture ( 15-16 ) with a more
strongly developed lip (13-14); finally,
they are taller at standardized whorl num-
bers during middle portions of ontogeny
(10-11). Thus, most of the information in
this large matrix reduces to a single contrast
between mottled (C. hendaUi) and ribby
(C ahacoense) morphotypes.

The third axis contains only 8.2 per cent
of the total information, but it also displays
a significant separation within the mottled
morphotype, C. bendaUi. \\\\\\ a single ex-
ception (sample 249, a peculiar, very large
and many-whorled, interior sample of Aba-
conian C. hendaUi), all samples with strong
loadings are from Grand Bahama. Factor
scores for this axis display a pattern of co-
variation found throughout the genus
(Gould et al., 1974; Gould and Paull, in
press ) : whorl number ( 3 ) and shell height
(7) are in negative association with mea-
sures of size at standardized whorl numbers.
Shells become large either by growing large
whorls (2, 10-11) or many whorls (3 and 7).

Shell height reflects whorl number because
shells add height but not width during later
growth; maximum width is reached early in
ontogeny in this genus named for a beehive.
If final size can vary only within narrow
limits, then these two alternate pathways
to a given size must covary negatively. The
primary geographic differentiation within
C. bendaUi on Little Bahama Bank has ap-
parently followed this common pattern of
covariance. Populations on Grand Bahama
have taken the route of small whorls leading
to high shells and many whorls ( high scores
for whorl number and shell height are
matched by high loadings of the same sign
for Grand Bahamian samples — Table 2 and
Figs. 6-7). Abaconian samples reach the
same sizes with fewer, larger whorls.

C) Coherence of Regional and Local Pat-
terns of Variation Within Morphotypes

Our consistent discovery of coherent,
broadly regional patterns of variation pro-
vides the primary datum for our rejection of
the traditional view about Cerion — that its
geographic variation is a "crazy-c[uilt"
formed by haphazard shifting about of
hundreds of species via hurricanes. We
haxe never failed to detect a hierarchy of
geographic coherence:

i) broad contiguous regions including
several islands have distincti\'e morphol-
ogies. C. striateUuni, the only Cerion
throughout the eastern regions of its range
( Hispaniola to the Virgin Islands ) , exhibits
a clinal pattern cjf variation with increas-
ing departure from "normal" morphology
away from major centers of distribution in
Cuba and the Bahamas (Gould and Paull,
1977). Cerion uva, the only species on
the outlying islands of Aruba, Bonaire, and
Curasao, is sufficiently distinct to warrant
its own subgenus in the traditional classifi-
cation (Pilsbry, 1902).

ii) islands within broad regions are un-
ambiguously, if subtly, distinct. The most
important discriminator of eastern cerions,
the first canonical axis of 23 samples, un-

388 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

covers the clinal pattern reported above
(Gould and Paull, 1977), but subsequent
axes clearly sort each island from all others
with no overlap. Ceriom of Aruba, Bonaire,
and Curayao also cluster by island ( Baker,
1924; Gould, 1969).

iii) contiguous geographic subregions
within islands can also be identified by very
minor, but thoroughly consistent, jjatterns
of character means and covariation; the
more isolated the subregion, the more dis-
tinct the morphology. The narrow "waist"
of Curasao, for example, separates popula-
tions of C. uva into two distinct groups
(Gould, 1969).

We will not venture any speculation
about adaptive values, importance of found-
ers, etc., but it does seem clear that geo-
graphic isolation is the primary correlate of
morphological variation within taxa of Ce-
rion. These patterns of geographic varia-
tion, by their stability and coherence, also
indicate that episodes of transport and
colonization have been rather less frequent
than tradition dictates.

The geographic variation of Little Ba-
hama Bank cerions conforms fully with
these new expectations of coherence. We
confine our comments to the mottled mor-
photype, C. hendalli since regional patterns
have never been demonstrated within it be-
fore. We do not have enough samples of C.
abacoense, and we have not seen two of its
three major populations in the field — Mores
Island and the apparently extinct popula-
tion of Grand Bahama. Nonetheless, tradi-
tional taxonomy has already recognized the
geographic coherence of four areas — Grand
Bahama, Mores Island, southern tip of
Abaco, and southeastern coast of Abaco.
We reject the names, but confirm the dis-
tinction in our morphometric analysis.

1. Separation of C. bendaUi from Grand
Bahama and Abaco. Figure 6 demonstrates
the morphological distinction of the two
islands (see discussion above). We are
particularly pleased to note that the basis
of separation is not a few static adult fea-
tures of unknown significance, but alternate

Table 3. Factor scores of original variables

UPON THE first Q-MODE AXIS FOR C. bendolH FROM

Grand Bahama.

1. protoconch width

0.249

2. 4th whorl width

0.289

3. total whods

0.188

4. 4th ribs

-0.129

5. 6th ribs

-0.077

6. 1st ribs

-0.075

7. height

0.259

8. width

0.295

9. protoconch height

0.189

10. 4th height

0.178

11. 4th-6th height

0.245

12. umbilical width

0.255

13. lip width

0.212

14. lip thickness

0.181

15. aperture height

0.282

16. aperture width

0.311

17. protrusion

0.201

18. tilt

-0.051

19. weight

0.240

20. aperture-suture

0.301

pathways of a major pattern in covariance
found throughout the genus.

2. The regional pattern on Grand Ba-
hama. As we collected on Grand Bahama,
it seemed to us that patterns of morphology
followed general trends throughout the
island. Shells of northern samples were
small, particularly in coastal populations
near mangrove areas. (This is another con-
sistent pattern within the mottled morpho-
type. Mottled shells are also dwarfed on
the low, bank-interior western coasts of
Andros, Eleuthera and Great Exuma).

We used trend surface analysis to test
a hypothesis of simple regional patterns.
This technique widely employed by geolo-
gists but little known among biologists
(Marcus and Vandermeer, 1966), performs
a multiple regression analysis of a mor-
phological feature (dependent variable)
against independent variables expressed as
geographic coordinates. Increasingly more
complex surfaces are obtained by adding
terms in a polynomial expansion of the X
and Y coordinates. Predictions from the best
fit surface are compared with actual values
to generate a vector of residuals that defines

Natural History Cerion VIII: A Revision • Gould and Woodruff 389

""Z.--'''' ^-rniy^

— .32
-- .38
.44

Figure 8. Third order trend surface analysis (with interaction terms suppressed) for projection of Grand Ba-
hamian samples on the first varimax axis of a Q-mode analysis; this is a "size" axis based on all variables.
Note simple pattern of increasing size from north to south, with more rapid transition near the northern coast,
where dwarfed samples pass rapidly to interior samples of modest size. Actual localities indicated by crosses.
This smooth variability has, in the past, been parcelled among three separate species defined only by differ-
ences in size.

"goodness of fit." The "art" of trend surface
analysis involves the selection of a fit that
explains enough information, yet remains
sufficiently simple to represent a truly re-
gional pattern. Points can be fit exactly
with polynomial surfaces of sufficiently
high order. We used the program of Lee
(1969).

We decided not to use the mean of in-
dividual characters as dependent variables,
but a value expressing major determinants of
covariance among samples. Consequently,
we performed a Q-mode factor analysis of
all C. bendaJU samples from Grand Bahama
and used loadings on the first varimax axis
(for a three-axis solution) as the dependent
variable. This single axis encompasses 57.2
per cent of the variance among 20 characters
for the 12 samples. Factor scores of vari-
ables upon it ( Table 3 ) show that it repre-
sents a fairly "pure" size axis, with high and
similar loadings for measures of final size
and whorl size. ( We do not detect the com-
mon negative interaction here, because we
do not consider alternate pathways to a

similar final size. We have, instead, the
opposite situation — a wide range of mean
shell size from very small on the north
coast to quite large elsewhere. The dwarfed
shells have both few whorls and small
whorls. )

The first order fit alone has a multiple
correlation of .82 for a coefficient of deter-
mination, r- = .67. A simple sloping plane
encompasses % of all geographic variation
expressed by the most important single di-
mension based on all 20 measured charac-
ters. As expected, the axis runs almost due
E-W with smaller values to the north. Fig-
ure 8 represents our highest surface, a third
order fit with interaction terms suppressed
(XiXo, Xi- Xo, and XiXo-— we did not have
enough sample points for the degrees of
freedom needed to fit them). This surface
yields a multiple correlation of .934, for a
coefficient of determination, r- = .87. Even
at this level of potential complexity, the
surface represents a surprisingly smooth
cline from small northern shells to larger
southern shells. Contour lines follow the

390 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

.3r

.2 -

0 -

-. I

-Fossil

Treasure
Cay

.5 .6

AXIS I

Figure 9. Factor loadings on the first and fourth axis
for all samples of Little Bahama Bank cerions. Al-
though the fourth axis explains less than 1 per cent
of the total variance among samples, it separates both
the Treasure Cay populations and the single fossil
sample from all others.

island itself, while bunching of lines at the
northern coast indicates tlie rapid transition
from coastal dwarfs to interior shells of
modest size that we observed in the field.
(Though we had noticed the coastal phe-
nomenon, we did not expect the regional
pattern to be so simple.) We detected no
geographic pattern in the vector of resid-
uals.

3. Distinction of subareas on Abaco. With
more than 30 samples of C. liendaUi from
Abaco, we could detect more local patterns
of distinction, also correlated with geo-
graphic isolation.

i) small, ribby shells of Pongo Carpet.
We have already reported in detail on a
semi-isolated coastal area well within the
range of C. hendalli (Gould, ^^'oodruff,
and Martin, 1974). Here, along nearly 7
km of eastern coast, we find a small, heavy,
fairly ribby morphology partly convergent
on C. ahacoewie. (We included only one
Pongo Carpet sample in this study; it has
the highest loading of any pure C. hendalli
sample upon the C. ahacoeme axis — Fig.
6. ) This Pongo Caipet morphology is most
distinct in its southern area of greatest
isolation, and varies in a clinal fashion to-
wards "normal" moiphology as it ap-
proaches the northern zone of contact. It
cannot be distinguished genetically from

surrounding populations of standard mor-
phology. In fact, it shares with these sur-
rounding normal populations the only dis-
tinctive genetic marker (the rare Mdh-2'''
allele) of its area — Mdh-2'' is fixed in all
other populations of C. hendalli. Although
these Pongo Carpet shells clearly merit spe-
cific distinction on all previous criteria, we
cannot regard them as any more than a local
variant within a coherent taxon.

ii) populations on Treasure Cay. The
difference between statistical and biological
significance is rarely appreciated. Mor-
phometricians routinely ignore axes of vari-
ation that encompass too little variation to
win statistically significant distinction from
zero. Yet minor patterns can be very real in
a biological sense. Suppose that we have
a large matrix with many samples and vari-
ables, and that a few samples from a geo-
graphically isolated region gain distinction
from all others by consistent differences in
just a few covarying characters. Suppose
also that this distinction is not evident in
qualitative observation. The information re-
corded by this distinction may include far
less than 1 per cent of the total matrix; yet
it is highly significant from a biological
point of view, especially since it is so easily
missed in raw data or qualitative observa-
tion. The criterion for importance must be
correlation with geography, not per cent of
information.

We offer such a case in the semi-isolated
samples of Treasure Cay (Fig. 3). The
fourth axis of cnu- Q-mode analysis for all
samples encompasses only 0.95 per cciit of
all information. Yet a plot of loadings upon
the fourth axis clearly separates all Treasure
Cay samples from all others with no overlap
(Fig. 9). Loadings for the Treasure Cay
samples never exceed 0.2, so the distinction
arises from less than 4 per cent of the infor-
mation (squared loading) in these popula-
tions. Factor scores for this axis (Table 4)
indicate that the separation of these sam-
ples arises from their high values for proto-
conch height and, to a lesser extent, whorl
number.

Natural History Cerion VIII: A Revision • Gould and Woodruff 391

iii) temporal variation. The carbonates of
Little Bahama Bank islands are largely
marine, and we do not find the soil zones
with abundant fossil cerions so common on
other islands. But we did collect one fossil
sample from an aeolianite in a cut on the
road leading to Snake Cay. We are espe-
cially pleased to report that this sample can
be distinguished clearly from all modern
populations, though its general appearance
links it unambiguously with living forms of
its area. As a strongly and fairly sparsely
ribbed sample of general C. bendaUi shape,
its mean morphology gives it an intermedi-
ate position in the essential distinction of
the two morphotypes (Fig. 6). Its unique-
ness is apparent in Fig. 8. It shares, with
Treasure Cay samples, the joint high values
of protoconch height and whorl number
(in fact, its loading on the fourth axis is
maximal among all samples), but it differs
from them in its weaker loading on the first
axis (i.e., its greater affinity with the ribby
morphotype ) .

The study of fossil cerions is yielding im-
portant information on the stability of mod-
ern patterns of geographic \'ariation within
taxa. In all three cases studied so far, fossil
samples share the same basic morphology of
modern populations, but the fossils occupy
presently unrealized portions of the mor-
phological spectrum (C. rude of St. Croix
vs. all living eastern cerions, Gould and
Paull, 1977; C. iiva from Indian shell mid-
dens on Cura9ao, Gould, 1971; and this
Snake Cay Road sample ) .

A note on technique: A potentially valid
objection has been raised against much
work in multivariate moiphometrics : avail-
able techniques for separation are now so
numerous and \'aried that proper selection
may be able to affirm nearly any a priori
preference. Robust conclusions may require
the joint confirmation of several techniques.
Readers may criticize our distinctions by
pointing to unusual features of our factor
analytic model; we use a Q-technique in
I-space while most workers prefer more
conventional R-mode analysis in A-space

Table 4. Factor scores for the 4th Q-mode
axis to illustrate the basis of distinction' ( in

COVARIAXCE) FOR THE TREASURE CaY SAMPLES.

1. protoconch width

-0.131

2. 4th whorl width

-0.243

3. total whorls

0.477

4. 4th ribs

-0.097

5. 6th ribs

-0.012

6. 1st ribs

-0.131

7. height

0.118

8. width

0.015

9. protoconch height

0.735

10. 4th height

0.036

11. 4th-6th height

-0.212

12. umbilical width

0.044

13. lip width

0.089

14. lip thickness

-0.011

15. aperture height

-0.084

16. aperture width

-0.068

17. protrusion

-0.159

18. tilt

-0.090

19. weight

-0.114

20. aperture-suture

-0.026

(Sneath and Sokal, 1973, p. 116). We nor-
malize vectors to eliminate size explicitly,
while most studies include these differences.
Finally, we equalize weights of variables
with an uncommon transformation, while
most studies use raw data or transform with
different techniques. Consequently, we re-
did the analysis in the R-mode with no nor-
malization or character weighting (using
BMD program P4M).

The factor loadings ( Table 5 ) display the
same pattern as the factor scores of our
Q-mode analysis with two interesting excep-
tions, one expected. The first axis of the R-
mode analysis reflects shell size, the vari-
ation explicitly eliminated in our Q-mode
analysis. The fourth axis displays a pattern
of covariance often seen in Cerion (Gould
et ah, 1974), but not encountered in our
Q-mode analysis. We find joint high load-
ings for four variables : apertural protrusion
and tilt (17-18) and lip width and thick-
ness ( 13-14 ) . When we specified our mea-
sures before beginning this study, we se-
lected these as potentially correlated traits
expressing the intensity of changes in

392 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 5. Factor loadings of oiugixal variables for an R-mode analysis of all Little Bahama

Bank samples.

1 . protoconch width

0.789

0.441

0.016

-0.021

0.190

2. 4th whorl width

0.931

-0.037

0.226

0.013

0.018

3. total whorls

0.329

0.288

-0.864

0.074

0.108

4. 4th ribs

-0.150

0.943

-0.195

0.083

-0.071

5. 6th ribs

-0.085

0.927

-0.268

-0.006

-0.048

6. 1st ribs

-0.144

0.918

-0.172

0.087

-0.078

7. height

0.900

0.000

-0.332

0.189

0.136

8. width

0.941

-0.251

0.015

0.033

0.099

9. protoconch height

0.200

-0.120

0.017

0.043

0.955

10. 4th height

0.305

-0.302

0.698

0.272

0.388

11. 4th-fith height

0.541

-0.412

0.709

0.102

-0.018

12. umbilical width

0.876

-0.269

-0.051

0.150

0.038

13. lip width

0.565

-0.536

0.001

0.407

0.164

14. lip thickness

0.485

-0.437

0.202

0.593

0.166

15. aperture height

0.909

-0.167

0.021

0.328

0.076

16. aperture width

0.910

-0.162

0.062

0.300

0.138

17. protrusion

0.689

-0.071

0.020

0.648

-0.018

18. tilt

0.090

0.407

0.019

0.809

0.017

19. weight

0.822

-0.318

0.113

0.299

0.072

20. aperture-suture

0.849

-0.070

0.135

0.023

0.147

growth that mark secretion of the adult
aperture (intense change in coiHng direc-
tion should be associated with a stronger
lip ) . We are gratified to see their joint asso-
ciation on an axis mathematically inde-
pendent of shell size. It would be less en-
lightening to find that intense development
correlated only with large shell size. The
association of size and adult development
exists to be sure ( 13, 14, and 17 also load
highly on the size axis), but the fourth
axis displays the partial independence of
adult development.

The other axes are essentially identical
with the factor scores of our Q-mode anal-
ysis. Axis 2 reflects the differences in rib-
bing that produced the basic separation
of mottled and smooth morphotypes in our
Q-mode analysis (axis 1); axis 3 records
the negative association of whorl number
and shell height with measures of whorl
size that separated Grand Bahamian and
Abaconian C. hendalli in our Q-mode anal-
ysis (axis 3); finally, axis 5 makes the same
separation of the Treasure Cay and Snake
Cay Road fossil samples from all others,
primarily on the basis of protoconch height.

When we consider factor scores to see
how these R-mode axes sort samples, we
find virtual identity with our loadings of
Q-mode analysis. The first axis is different,
since we eliminate its effects by normaliza-
tion of sample vectors in our Q-mode anal-
ysis. The R-mode first axis merely sorts
samples by shell size — a biologically un-
enlightening distinction in this case. But
axes 2, 3, and 5 make the same separations as
corresponding axes in the Q-mode analysis.
The correlation coefficients ( at N = 52 ) for
R-mode scores with Q-mode loadings for
corresponding axes are .87 for R-mode 2
with Q-mode 1 (ribbing) to separate the
moiphotypes; .60 for R-mode 3 with
Q-mode 3 ( negative interaction of whorl
number and whorl size to separate Grand
Bahama and Abaco C. hendalli); .72 for
R-mode 5 with Q-mode 4 (to separate
Treasure Cay and Snake Cay Road fossil
samples from all others). We are therefore
confident that our Q-mode patterns identify
real and important distinctions in nature,
robust with respect to techniques used to
identify them, and not artifacts of unusual
multivariate procedures.

Natural History Cerion VIII: A Revision • Gould and Woodruff 393

Table 6. Univariate ANOVA for discrimintatory power of original variables in samples of the
hybrid zone at rocky point. univariate f -ratios with 4 and 89 degrees of freedom.

Variable

among mean sq.

within mean sq.

F-ratio

Probability

1. protoconch width

23.33

9.59

2.43

0.5242E-01

2. 4th whorl width

136.52

20.35

6.71

0.2163E-03

3. total whorls

0.25

0.14

1.83

0.1299E+00

4. 4th ribs

1046.82

32.45

32.26

0.1014E-07

5. 6th ribs

608.14

18.56

32.77

0.9304E-08

6. 1st ribs

18.53

2.76

6.72

0.2131E-03

7. height

36.94

2.80

13.22

0.2309E-05

8. width

5.87

0.32

18.27

0.2895E-06

9. protoconch height

5.64

6.46

0.87

0.5149E+00

10. 4th height

16.83

13.50

1.25

0.2963E+00

11. 4th-6th height

3.84

0.99

3.89

0.6083E-02

12. umbilical width

265.17

29.25

9.07

0.2865E-04

13. lip width

59.04

18.88

3.13

0.1843E-01

14. lip thickness

85.70

19.15

4.47

0.2774E-02

15. aperture height

465.77

28.87

16.13

0.6345E-06

16. aperture width

166.91

20.18

8.27

0.5324E-04

17. protrusion

32.23

10.79

2.99

0.2271E-01

18. tilt

14.48

8.33

1.74

0.1474E-f00

19. weight

2.04

0.05

40.03

0.3185E-08

20. aperture-suture

169.07

27.66

6.11

0.3978E-03

D) Interaction Between Morphotypes

We recorded the areas of interaction be-
tween ribby (C. ahacoense) and mottled
(C. hendaUi) morphotypes in our discussion
of geographic distribution (pp. 376-377).
We identified these areas before perfonning
any multivariate analysis upon the shells.
Thus, the intermediate position of all these
samples in the contrast between ribby and
mottled morphotypes (axes 1 and 2 of our
Q-mode analysis) serves as a strong con-
firmation of interaction. The intermediate
field of Figure 6 is shared by only two other
samples: the convergent Pongo Carpet
sample ( No. 316 ) lies on the border of mot-
tled and intermediate samples; secondly, the
fossil sample (No. 246) occupies an inter-
mediate position.

All other points in the intermediate zone
belong to samples in areas of geographic
contact between the morphotypes. These
include :

1. the sample from Sandy Point Village
(Fig. 6) marking the transition from exte-
rior to interior coast (sample 251).

2. samples from the main road at The

Crossing (Fig. 6 — samples 253 and 255)
where interior C. bendalli is separated by
500 m and a narrow hill from coastal C
ahacoense. We did not record these in the
field as intermediate in morphology; they
appeared to us at the time as somewhat
peculiar C. bendalli. Their intermediate
position on Figure 6 indicates that some
leakage occurs in this area of closest geo-
graphic contact between morphologies ap-
parently separate in the field.

3. the hybrid zone at Rocky Point. We
noted in the field that the transition from
southern ribby to northern mottled seems to
occur in the narrow area between samples
305-260. We are therefore pleased to dem-
onstrate a smooth transition in morphology,
spanning the entire range from pure ribby
to pure mottled, along the geographic se-
quence in this area: 305-306-307-308-260
( Fig. 6 ) . Our impression that the effects of
hybridization do not spread far south of 305
seems to be affirmed by the non-clinal posi-
tions of the next two southern samples, 304
and 259. Finally, the interior samples of C.
l)endaUi collected in the area of coastal hy-

394 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 7. Matrix of Mahalanobis D" distances
for samples of the hybrid zone at rocky point.

305

306

307

308

260

305 0.0

306 8.81493 0.0

307 19.71928 6.93176 0.0

308 27.42574 14.81611 5.17647 0.0

260 36.26627 19.43301 10.82527 4.94755 0.0

bridization confirm the localization of in-
teraction. Sample 310, collected 400 m from
the coast between sample 305 (the pure
ribby beginning of the hybrid zone) and
sample 306 lies among mottled samples, but
near the peripheiy of mottled and inter-
mediate forms. Sample 309, about 600 m
inland from 310, is well within the C. ben-
dalli cluster and shows no signs of inter-
mediacy.

We then performed a discriminant anal-
ysis on samples of the hybrid zone, using
D/DA, a program written by John Rhoads,
Dept. of Anthropology, Yale University ( see
Gould et al., 1974 for more details). In the
field, we had concluded that the morpho-
logical effects of hybridization are confined
to a small, coastal area at Rocky Point (Fig.
5). We therefore performed our analysis
on the five samples collected along this mile
of coast (from south to north, 305, 306, 307,
308, and 260).

The table of univariate ANOVA's ( Table
6) shows that the best discriminators are
measures of ribbing and shell size — scarcely
surprising .since shells of C. hendalli are
characteristically smaller and more copi-
ously (though more weakly) ribbed than
those of C. ahacoense.

As a first indication of evenly clinal pat-
terns, the matrix of Mahalanobis D- dis-
tances (an overall measiue of similarity
based on all characters with variance and
covariance adjustments) exhibits a smootli
morphological transition along the geo-
graphic axis of collections (Table 7). Fig-
ure 10 represents a plot of all samples
against the first two discriminant axes. The
first axis, which encompasses fully % (74.3
per cent) of all information, arrays the sam-

Figure 10. Samples from the hybrid zone at Rocky
Point projected on the first two axes of a discriminant
analysis. Points represent an analysis based only on
samples that, from our field impressions, constitute the
zone of transition (numbers 305-306-307-308-260 from
south to north). Note the smooth transition along the
first axis (74.3 per cent of ail information). Crosses
represent a separate analysis (shown here on the
same scales) for these samples plus two more
southern samples in the same area (304 and 259).
Sample 304 breaks the morphological dine, thus con-
firming our impression that it is beyond the localized
zone of interaction.

pies in a smooth and gradual transition.
The second axis (only 13.2 per cent) pro-
duces the "horseshoe pattern" so commonly
seen when two axes exhaust nearly all infor-
mation (Reyment, 1975). (If end-member
samples have high values on the main dis-
criminator and intermediate samples lie
close to zero, then the second axis must
emphasize these intermediate samples.)

The table of discriminant loadings dis-
plays the patterns of covariance that sepa-
rate samples (Table 8 — these loadings are
correlations of original variables with new
axes, not coefficients of the discriminant
axes themselves). Ribbing (positive load-
ings) and adult size (negative loadings)
dominate the first axis. Northern (C. hen-
dalli) samples with their numerous ribs and
small shells have high positive projections
upon this axis. Moving southward through
the hybrid zone, shells gradually become
larger as ribs become sparser and stronger.
(Measures of ribbing and size are invari-
ably independent as patterns of covariance
within samples — see Gould et al., 1974.
They are united as joint discriminators of
the morphotypes in this study of among
sample covariance.) Loadings on the mi-

Natural History Cerion VIII: A Revision • Gould and Woodruff 395

Table 8. Discriminant loadings of original
variables upon axes used to separate samples

OF THE HYBRID ZONE AT RoCKY PoiNT.

Table 9. Classification (hits and misses) ta-
ble FOR discriminant ANALYSIS OF SAMPLES FROM
THE HYBRID ZONE AT RoCKY PoiNT. TOTAL HITS =

79 OUT OF 94 POSSIBLE. Rate = .8404.

1. protoconch width

2. 4th whorl width

3. total whorls

4. 4th ribs

5. 6th ribs

6. 1st ribs

7. height

8. width

9. protoconch height

10. 4th height

11. 4th-6th height
umbilical width
lip width
lip thickness
aperture height
aperture width
protrusion
tilt

19. weight

20. aperture-suture

12.
13.
14.
15.
16.
17.
18.

-0.0377
-0.2121
-0.0300
0.5185
0.5322
0.2313
-0.3043
-0.3785
-0.0767
-0.0965
-0.1753
-0.2312
-0.1008
-0.1716
-0.3355
-0.2413
-0.1074
0.1210
-0.5706
-0.1840

0.1390
0.0888

-0.1247
0.3257
0.2315
0.1053
0.0137

-0.0351

-0.0508
0.0967
0.0695
0.2911

-0.1465

-0.0549
0.2920

-0.0571
0.2179
0.0371
0.1420

-0.3474

nor, second axis make little biological sense
to us; they seem to represent a concatena-
tion of those variables that distinguish, in a
minor way, the intermediate samples from
both endpoints. Thus, any measure that
distinguishes either endpoint has a rela-
tively high loading ( ribs and size now have
joint positive loadings), while three dis-
parate measures with generally higher val-
ues in the intermediate samples (variables
3, 13, and 20) have negative loadings.

As a final example of smooth transition.
Table 9 presents a matrix of classification.
Seventy-nine of 94 specimens lie nearest to
their own sample centroids (84 per cent).
Every misclassified individual groups with
a geographically adjacent sample.

This smooth transition is matched by a
total lack of evidence for any increased
variability in intermediate samples (as we
might expect in a "classic" hybrid zone —
Mayr, 1963). Table 10 presents C.V.'s for
all 5 samples and for typical samples of
"pure" mottled and ribby shells in the cen-
tral areas of their distribution.

In an attempt to learn whether the

305

306

307

308

260

305

18.

306

15.

307

17.

308

11.

260

18.

smooth morphological transition continues
southward beyond our perception of it in
the field, we performed a similar analysis
on 7 samples — the 5 used before plus 304
and 259, the next southern coastal samples
(Fig. 10). The first axis is virtually un-
changed in both loadings and discrimina-
tory power. As the previous factor analysis
indicated ( Fig. 6), the next southern sample
( 304 ) breaks the smooth transition by plot-
ting closer to the C. bendaUi axis than sam-
ple 305 directly to the north.

This morphometric analysis cannot re-
solve the key question of appropriate
biological status for populations of the two
morphotypes. Are they imperfectly sepa-
rated entities meriting taxonomic recogni-
tion as semispecies or simple geographic
variants with uneventful and unrestricted
mixture at points of contact? The habitat
preferences and coherence in areas of near
contact (The Crossing) might argue for
separation, and the very localized nature of
the hybrid zone would support such an
assertion. But rapid transitions and step
clines occur within coherent taxa (Endler,
1977 ) . We must turn to genetic analysis for
further enlightenment.

IV. ALLOZYME VARIATION OF CERION
ON THE LITTLE BAHAMA BANK

A. Introduction

The practice of combining electrophoresis
with histochemical staining methods to
study variation of enzymes is now well
established. The applicability of this meth-

396 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 10. Coefficients of variation for hybrid zone samples and for representative samples
OF C. bendalli and C. abacoense from the center of their ranges

central

bendalli
214

„ hybrid zone -

abacoense

260

308

307

306

305

250

1. protoconch width

6.3

4.7

5.7

5.9

6.2

4.8

5.2

2. width of 4th whorl

5.4

5.2

4.6

7.3

7.1

4.8

6.1

3. total whorls

4.9

4.7

5.1

5.0

5.2

4.7

5.6

4. ribs fourth whorl

19.5

10.8

12.9

11.2

11.6

12.0

11.0

5. rilis sixth whorl

20.1

10.4

11.9

10.4

8.6

6.7

11.3

6. first ribs

22.3

12.0

11.8

13.8

16.6

18.4

19.4

7. shell length

8.0

6.2

6.9

4.5

6.3

6.8

8. shell width

4.1

3.6

4.9

5.0

5.8

4.4

4.3

9. protoconch height

14.0

9.4

9.5

10.5

12.3

10.5

8.8

10. height 4th whorl

5.4

5.2

6.3

9.0

6.9

6.3

8.0

11. height 4-6

7.6

9.3

9.4

9.8

11.3

10.3

8.3

12. umbilical width

8.0

12.3

13.1

11.6

12.0

9.9

13. lip width

15.3

14.9

27.0

19.3

22.2

23.0

14.4

14. lip thickness

31.1

32.5

18.8

24.7

26.0

26.6

24.1

15. aperture height

7.4

7.2

5.0

5.7

7.1

5.3

3.9

16. aperture width

6.5

5.8

4.9

6.9

6.4

5.5

17. protrusion

16.7

14.2

12.9

16.5

13.0

12.7

12.3

18. tilt

16.4

16.5

20.0

10.7

13.3

12.9

17.9

19. weight

27.0

25.8

14.2

15.3

14.1

15.3

13.9

20. aperture to suture

9.2

8.5

17.5

7.9

12.7

9.0

11.6

odology to current problems of evolutionary
biology is well reviewed by Avise ( 1974 ) ,
Lewontin ( 1974 ) , and various authors in
the volume edited by Ayala (1976). We
originally appHed this approach to Cerion
in the hope that genie variability might be
more conservative than shell form in these
morphologically variable animals. In the
following account, we will report our find-
ings as they apply to the systematic problem
of the relationship between ribby and mot-
tled morphotypes. For convenience, these
contrasting shell types will be referred to C.
ahacoen.se and C. JyendaUi respectively as
suggested above. In subsequent papers, we
will describe the genetic aspects of the in-
teraction between these taxa in more detail
( Woodruff and Gould, in press ) , and the
relation between genie and phenic variation
at the level of the individual, population,
and species (Woodruff, in prep.).

\'ariation in the electrophoretic pattern
of structural gene products was surveyed in
1,575 individual adult snails from 47 pop-
ulations from the islands of the Little Ba-

hama Bank. Localities are indicated in
Figure 3. In most cases, these are the
same localities described in the morpho-
metric analyses presented above. Further-
more, whenever possible, we have examined
the same individual snails. Sample prepara-
tion, biochemical specifics, and other tech-
nical aspects of the starch gel electro-
phoresis apparatus employed are described
elsewhere (Woodruff, 1975b). In the con-
text of this survey, we have examined 16
enzyme systems and some general proteins
and interpreted the observed banding pat-
terns in terms of at least 28 loci. Here we
will describe the variation in 20 of these
structural gene products: ones that we
found to give reproducible and genetically
interpretable patterns. Variation of these
enzymes among the Little Bahama Bank
Cerion is outlined in Table 11. While 14
of these proteins are monomorphic and are
fixed for the same allele in both taxa,
polymorphisms were detected in the re-
maining six. (Est-3 and Pgi are also vari-
able allozymes but are excluded from this

Natural History Cerion VIII: A Revision • Gould and Woodruff 397

discussion. ) Variation in each case is due to
a simple Mendelian system involving co-
dominant alleles. In the absence of formal
genetic crosses, our genetic interpretations
are based on two criteria. First, phenotypic
ratios (and presumed genotypic ratios)
agree closely with Hardy-Weinberg expec-
tations. Second, patterns of banding of
particular enzymes correspond to simple
models of molecular structure. In most
cases, the inferred structures are similar to
those of functionally analogous enzymes in
other animals whose structures have been
established by other techniques. The six
polymorphic loci segregate independently
of one another; this is quite reasonable as
one species of Cerion is known to have 27
pairs of chromosomes (Burch and Kim,
1962).

Before proceeding with the results of this
genetic survey, we wish to outline the gen-
eral nature of population structure in Ce-
rion. Beginning in 1973, Woodruff has
been studying two large marked popula-
tions of C. bendaUi and C. abacoen.se on
Abaco (3 years' experience with over 1,500
individually marked snails). Generation
time for Cerion is not well defined. Juve-
niles grow slowly and erratically and prob-
ably do not lay down the shell's adult lip
until they are 3 years old. The duration of
the adult phase is also poorly defined;
multiple-recapture studies suggest that
some adults live at least another 10 years.
Snail distribution at the coastal study sites
is patchy but averages 8-13 adults per
square meter. Dispersal data are now being
used to estimate various evolutionarily im-
portant parameters. Effective neighbor-
hood size or effective population size (N
of Wright, 1946) is about 1,000 snails.
Neighborhood area is 50-100 m-. Our
preliminary estimate for gene flow ( / of
May et al., 1975) suggests that this variable
will be shown to have a value of about 3
meters. This estimate is, however, based on
the formula / = xVg where x is the mean
distance travelled in a generation and g is
the probability of leaving a deme or neigh-

Table 11. Electrophoretically demonstrable

ALLOZYMIC VARIATION IN Ccriotl FROM THE LiTTLE

Bahama Bank.

No.
of
Enzyme alleles

7
*

Variable Enzymes:

Esterase-2 (Est-2)

Esterase-3 (Est-3)

Malate dehydrogenase- 1 (Mdh-1)

Malate dehydrogenase-2 (Mdh-2)

6-Phosphogluconate dehydrogenase-l

(6-Pgdh)
Phosphoglucose isomerase (Pgi)
Glutamic oxalacetic transaminase (Got-1) 3
Leucine aminopeptidase ( Lap ) 3
Invariable enzymes : alcohol dehydrogenase ( Adh ) ,
Est-1, Est-6, Est-7, isocitrate dehydrogenase-l
(Idh-1), Idh-2, a-glycerophosphate dehydrogenase
(a-Gpdh), indophenol oxidase (Ipo), phosphoglu-
comutase (Pgm-1), Pgm-2, Got-2, acid phospha-
tase-1 (Acp-1), Acp-2, alkaline phosphatase- 1
(Ap-1)

* At least two alleles segregating; variation not
yet interpretable.

borhood. While these parameters can be
estimated fairly accurately in Cerion, the
ultimate determinant of effective gene flow,
reproductive success outside the deme of
birth, will be very difficult to assess.

One of the first things we were able to
estabhsh was that Cerion, a facultative her-
maphrodite (Richter, 1926; Jaenicke, 1933),
is apparently outbreeding. This conclusion
is based on the close agreement between
observed and expected genotype frequen-
cies in all the larger (N > 30) samples.
This concordance is particularly impressive
in the samples where four and five alleles
are segregating at the Est-2 locus. In addi-
tion, estimates of inbreeding ( F of Wright,
1965) and outcrossing (X, where F =
(1 _ X/1 + X), Nei and Syakudo, 1958)
were also calculated for the four largest
samples. In each case, lambda was greater
than 0.96 thus confirming our conclusion
regarding panmixia with respect to the al-
lozymes studied.

The second important finding about Ce-
rion was that like most other organisms

398 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 12. Variation in malate dehydrogenases and 6-phosphogluconate dehydrogenase.

Allele frequency

Number of

Locality snaUs Mdh-l" Mdh-1'' Mdh-2-' Mdh-2''

Grand Bahama — C. hendalli

211 70 0.40 0.60 ...... 1.00

208 1 1.00 .___ -....- 1.00

204B 64 0.65 0.35 ...... 1.00

205 38 0.49 0.51 . 1.00

207 6 0.50 0.50 ._-_. 1.00

202 38 0.51 0.49 -.... 1.00

200 6 0.42 0.58 . 1.00

201 11 0.50 0.50 - 1.00

Little Ahaco — C. hendalli

213 35 0.20 0.80 ...... 1.00

214 37 0.27 0.73 ..-.. 1.00

216 36 0.18 0.82 0.14 0.86

Great Abaco — C. hendalli

217 18 0.28 0.72 0.08 0.92

218 38 0.26 0.74 0.13 0.87
224 45 0.32 0.68 0.01 0.99
223 25 0.36 0.64 0.02 0.98
222 39 0.35 0.65 0.03 0.97
226 53 0.26 0.74 0.01 0.99
220 56 0.25 0.75 ..... 1.00

219 77 0.36 0.64 _.-__. 1.00
316 14 0.43 0.57 .-... 1.00
228 21 0.36 0.64 0.05 0.95

230 7 0.29 0.71 .... 1.00

231 15 0.27 0.73 0.03 0.97
240 37 0.28 0.72 .._... 1.00
247 37 0.34 0.66 ...... 1.00

243 102 0.29 0.71 -_.... 1.00
245 42 0.31 0.69 _._... 1.00

244 41 0.37 0.63 _.. 1.00

Great Abaco — area of interaction between C. hendalli and C. ahaeoense

260 36 0.28 0.72 ...... 1.00

501 12 0.33 0.67 ...... 1.00

308 24 0.21 0.79 ...... 1.00

307 24 0.25 0.75 ...__. 1.00

306 36 0.31 0.69 ._.__. 1.00

305 29 0.34 0.66 1.00

304 36 0.29 0.71 . 1.00

311 24 0.27 0.73 ...... 1.00

253 36 0.39 0.61 ...... 1.00

255 30 0.35 0.65 1.00

251 84 0.43 0.57 . 1.00

Great Abaco — C. ahaeoense

259 13 0.25 0.75 . 1.00

520 7 0.36 0.64 ...... 1.00

521 36 0.28 0.72 ...... 1.00

254 34 0.29 0.71 ...... 1.00

257 12 0.42 0.58 ..... 1.00

252 24 0.25 0.75 ...... 1.00

250 70 0.36 0.64 1.00

6-Pgdh'>

e-Pgdhb

0.64

0.36

1.00

0.54

0.46

0.67

0.33

0.83

0.17

0.64

0.36

0.67

0.33

0.68

0.32

1.00

0.90

0.10

1.00

0.81

0.19

0.69

0.31

0.57

0.43

0.53

0.47

0.60

0.40

0.56

0.44

1.00

0.62

0.38

0.93

0.07

0.40

0.60

0.78

0.22

0.75

0.25

0.79

0.21

1.00

Natural History Cer/o.v VIII: A Revision • Gould and Woodruff 399

Table 13. Variatiox in glutamic oxalacetic transaminase axd leucine aminopeptidase.

Locality

Number of
Snails

Got-l«

Got- lb

Allele frequency

Got-1'^

Lap-1"

Lap-l"

Lap-1'^

Grand Bahama —

C. bendalli

211

0.67

0.33

0.73

0.14

0.13

208

1.00

204B

0.77

0.23

0.74

0.24

0.02

205

0.84

0.16

0.78

0.17

0.05

207

0.33

0.67

0.92

0.08

202

0.55

0.45

0.89

0.11

200

0.58

0.42

0.92

0.08

201

0.47

0.53

0.77

0.23

Little Abaco — C.

bendalli

213

0.60

0.40

0.81

0.19

214

0.66

0.34

0.86

0.14

216

0.43

0.57

0.96

0.04

Great Aliaco — C

bendalli

217

0.56

0.44

0.97

0.03

218

0.54

0.46

0.95

0.05

224

0.51

0.49

0.96

0.04

223

0.56

0.44

0.96

0.04

222

0.59

0.41

0.95

0.05

226

0.48

0.52

0.96

0.04

220

0.46

0.54

0.95

0.05

219

0.51

0.49

0.99

0.01

316

0.54

0.46

0.89

0.11

228

0.50

1.00

230

0.57

0.43

1.00

231

0.53

0.47

0.97

0.03

240

0.55

0.45

0.96

0.04

247

0.58

0.42

0.93

0.07

243

102

0.47

0.53

0.87

0.13

245

0.54

0.46

0.90

0.10

244

0.52

0.48

0.95

0.05

Great Abaco — area of interaction between C

bendalli

and C. abacoense

260

0.58

0.42

0.90

0.10

501

0.54

0.46

0.88

0.12

308

0.50

0.42

.08

0.94

0.06

307

0.69

0.31

0.81

0.19

306

0.56

0.44

0.78

0.22

305

0.57

0.43

0.90

0.10

304

0.60

0.40

0.86

0.14

311

0.44

0.54

0.02

0.90

0.10

253

0.53

0.47

0.67

0.14

0.19

255

0.48

0.50

0.02

0.72

0.13

0.15

251

0.51

0.47

0.02

0.83

0.04

0.13

Great Abaco — C

. abacoense

259

0.62

0.38

0.81

0.19

520

0.36

0.64

0.50

521

0.32

0.68

0.82

0.18

254

0.28

0.72

0.73

0.09

0.18

257

0.58

0.42

0.79

0.13

0.08

252

0.38

0.62

0.77

0.23

250

0.61

0.38

0.01

0.71

0.22

0.06

400 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 14. Variation in esterase-2.

Allele frequency

Number of ■ — — —

Locality Snails Est-2« £51-2" Est-2<> £51-2" Est-2« Est-2' Est-28

Grand Bahama — C. bcndalli

211

208

204B

205

207

202

200

201

Little Abaco — C. bendaUi

213 35

214 37 0.01 0.01

216 36

Great Abaco — C. bendalli

217 18

218 38
224 45
223 25
222 39
226 53
220 56

219 77
316 14
228 21

230 7

231 15 . 0.03

240 37

247 37

243 102
245 42

244 41

260 36 __.... 0.03

501 12 0.04

308 24 __-... 0.02

307 24 . 0.08

306 36

305 29 _..... 0.02

304 36 0.03

311 24 ...... 0.08

253 36 ...... 0.09

255 30 ...... 0.10

251 84

Great Abaco — C. abacoense

259 13

520 7

521 36

254 34

257 12 . 0.04

252 24
250 70

0.04

0.18

0.24
1.00

0.54

<0.01

0.05

0.14

0.46

0.32

0.03

0.07

0.54

0.33

0.03

0.08

0.42

0.08

0.01

0.20

0.14

0.58

0.07

0.17

0.83

0.23

0.09

0.68

0.10
0.22

0.90
0.76
1.00

0.22
0.04

0.78
0.96

0.06

0.94

0.10

0.90

0.03

0.97

0.07

0.93

0.02

0.98

0.05

0.95

0.04
0.02
0.14
0.10

0.96
0.98
9.86
0.87
1.00
1.00

0.03

0.97
1.00
1.00

idalli

and C. abacoense

0.64
0.58
0.52

0.33
0.38
0.46

0.40
0.29
0.26
0.57

0.52
0.71
0.72
0.40

0.54
0.85

0.31
0.06

0.06

0.85
0.50

0.05
1.00

0.50
1.00

0.19

0.81

0.28
0.50

0.72
0.46

0.25

0.75

0.04

0.96

Natural History Cerion VIII: A Revision • Gould and Woodruff 401

Figure 11.
Figure 3.

Geographic variation in malate dehydrogenase-1. Sample locality numbers may be discerned from

with open population structures, it has a
rich endowment of genetic variabihty. The
proportion of loci that are polymorphic per
population (P) was in the range of 20-30
per cent ( 20-36 per cent if the variable but
uninterpretable Est-3 and Pgi loci are in-
cluded). Average heterozygosity per indi-
vidual (H) was in the range of 5-12 per
cent. Interpretation of variation in P and
H will be discussed below after the geo-
graphic variation in the polymorphic allo-
zymes has been described.

B. Geographic Variation in Polymorphic
Loci

Having established that C. bendalli and
C ahacoense were identical with respect to
14 genetic loci, we turned our attention to
variation at the six polymorphic loci. We
will present these data in two ways. In

Tables 12-14, the allele frequencies are
shown with the localities grouped according
to geographic and taxonomic constraints.
The decision as to whether a sample was
placed in category 4 (transition zone) in
the tables or in category 5 (C. ahacoense)
was based on shell morphology as outlined
in the previous section. In Figures 11-16,
the presentations are not biased by any a
priori taxonomic constraints. Note that the
smallest samples are not shown in the fig-
ures. We have initially resisted the tempta-
tion to group our samples according to
island or sample region in any more formal
sense, because the population structure of
these animals suggests that such data pool-
ing could seriously distort our conclusions.
Malate dehydrogenase-1. Two NAD de-
pendent alleles have been demonstrated at
this locus in C. bendalli from Abaco (Gould

402 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

0OOOOOQ

Mclh-2

o
O

o
o
o
o
o
o
o
o
o
o
o

Figure 12. Geographic variation in malate dehyclrogenase-2. Sample locality numbers may be discerned from
Figure 3.

ct al., 1974; \\'oodmff, 1975). Data in Ta-
ble 12 and Fig. 11 indicate that there is no
consistent difference in allele frequency
between C. bendalli and C. ahacoen.se
morphot\'pes on Great Abaco where
Mdh-T' varies in frequency between 0.21-
0.43 in both taxa. Elsewhere, frequencies
of Mdli-1" are slightly different: being
lower on Little Abaco (O.lS-0.27), and
higher on Grand Bahama (0.40-0.65). Al-
lele frequencies in adjacent populations are
similar, and no dramatic shifts or clines in
allele frequency were noted. There is no
marked change in this allozyme in the area
of interaction between C. bendalli and C.
ahacocnse on Great Abaco.

Malate deJuidrofienu.se-2. Gould et al.
( 1974 ) found two alleles at this locus in
populations of C. bendalli from northern

Great Abaco. NAD-Mdh-2^' is a rare allele
(0.01-0.04) that occurs in standard C. ben-
dalli and in some samples of the aberrant
Pongo Carpet morphotype found in this
area. This allele was detected in popula-
tions extending from the eastern end of
Little Abaco (Loc. 216) south to Treasure
Cay, a distance of about 30 km. (It was
not found in the three most isolated (and
morphometrically differentiated) of the
Pongo Carpet samples.) Subsequently,
Woodruff (1975) reported Mdh-2" was ab-
sent in 3 samples of C. bendalli from locali-
ties about 50 km. south of this area. We
now report that the Mdh-2'' allele has not
been detected in over 1,000 snails from else-
where on the Little Bahama Bank (Table
12, Fig. 12). We conclude that C. bendalli
and C. abacoense are not differentiated at

Natural History Cerion VIII: A Revision • Gould and Woodruff 403

Figure 13. Geographic variation in 6-phosphogluconate dehydrogenase. Sample locality numbers may be dis-
cerned from Figure 3.

this locus and that the presence of the rare
allele in northern Great Abaco is a bio-
chemical area effect. Until more is known
about the variation of this gene elsewhere
in the Bahamas, we are inclined to invoke
mutation and the spread of a locally ad-
vantageous allele as the most likely explana-
tion for this phenomenon. An alternative
hypothesis — that Mdh-2=' was introduced
into the area with the aberrant Pongo Car-
pet morphotype by hurricane transport
from elsewhere — is rejected at present be-
cause the allele was not detected in the
three most differentiated populations of the
Pongo Carpet snails.

6-Phospho gluconate dehydrogenase.
While populations of C. bendalli from near
Pongo Caipet (Gould et al., 1974) and
Snake Cay, Great Abaco (Woodruff, 1975)
are monomorphic for 6-Pgdh'\ a second
codominant allele (6-Pgdh-l'') has been

found elsewhere on the Little Bahama Bank
(Fig. 13). This allozyme stains as a single,
sharp band of slightly reduced mobility
relative to e-Pgdh-l-"*; heterozygotes are
3-banded. As shown in Table 11 and Fig.
13, all samples from Grand Bahama are
polymorphic with 6-Pgdh-T'' varying in fre-
quency between 0.54-0.83. On Little and
Great Abaco, the 6-Pgdh-l'' allele has been
found in two separate areas. First, at the
eastern end of Little Abaco (Loc. 216), 7
heterozygotes were noted among 36 snails
examined. No trace of this allele was found
in the sample from Loc. 217 which was
collected 100 m away on Great Abaco at
the other end of the causeway connecting
the two islands. Sixty kilometers further
south, the e-Pgdh-L' allele was found again
in samples from the area of interaction be-
tween C. bendalli and C. abacoense. It was
detected in 6 of 11 samples between Rocky

404 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Got-l

3
3
3

Figure 14. Geographic variation in glutamic oxalacetic transaminase. Sample locality numbers may be dis-
cerned from Figure 3.

Point and The Crossing and in 5 of 6 sam-
ples of C. ahacocnse morphotype. Its fre-
quency in tliis area ranged up to 0.60 in the
populations where it was detected. It was,
however, conspicuously absent in samples
of intermediate morphotype from the west
side of The Crossing (Locs. 253, 255) and
Sandy Point (Loc. 251) and in the "pure"
C. abacoeme from Hole-in-the-Wall ( Loc.
250). The isolated occurrence of 6-Pgdh-l''
on Little Abaco is tentatively interpreted as
being due to mutation and drift. The oc-
currence of this allele at higher frequencies
on Grand Bahama and in the transition zone
between the morphotypes on Great Abaco
must be due to other forces.

Glutamic oxalacetic transaminase. In C.
hemlalli, two equally common alleles were
found in 12 populations from northern and
central Great Abaco (Gould et al., 1974;
Woodruff, 1975). As seen in Table 13 and

Fig. 14, Got-l'' occurs at a slightly higher
frequency than Got-P' throughout the range
of this taxon. Got-l'' reaches its greatest
frequencies (0.77-0.84) in western and in-
terior samples from Grand Bahama. Seven
samples of the ribby morphotype, C. ah-
acoetise, were also examined and found to
have Got-l-' frequencies of 0.28-0.62. Fre-
quencies of 0.44-0.69 were found to char-
acterize samples from the area of interac-
tion between these taxa. A third allele,
Got-1*^ is now reported from southern Great
Abaco where it occurs at low frequency
( 0.01-0.08 ) . It has been detected in "pure"
C. abacoense (Loc. 250) and 4 samples of
intermediate morphotype from Sandy Point
(Loc. 251), The Crossing (Loc. 255) and
near Rocky Point (Locs. 308, 311). This
rare allele has yet to be found in "pure" C.
bendaUi from Abaco or Grand Bahama.
In mobility, Got-1^ is slower than Got-l'';

Natural History Cerion VIII: A Revision • Gould and Woodruff 405

Figure 15. Geographic variation in leucine aminopeptidase. Sample locality numbers may be discerned from
Figure 3.

like the latter the former stains a well-de-
fined doublet, the heterozygotes having
four bands.

Leucine aminopeptidase. We have pre-
viously shown that Lap-1'^ is the predomi-
nant allele in C. bendalli near Pongo Carpet
and Snake Cay (Gould et al., 1974; Wood-
ruff, 1975). In these populations a slower
allele (previously designated Lap-1'') oc-
curred at a frequency of up to 0.13. Now
we report finding a third allele of inter-
mediate mobility in Cerion from Grand Ba-
hama and southern Great Abaco (Table 13,
Fig. 15). For consistency, the newly dis-
covered allele is now designated Lap-1'^;
the designation of the slowest allele is ac-
cordingly changed to Lap-l''.

Lap-1=* is the common allele throughout
the Little Bahama Bank: it varies in fre-

quency and is typically over 0.85 (range:
0.67-1.00). Lap-1'^' is also widespread. Its
absence in a few samples is presumably due
to sampling error. No particular biological
significance is attached to the minor inter-
population variation in frequency of this
allele. The third allele, Lap-1'', is known
from 5 localities in western Grand Bahama
and from 6 localities at the southern end of
Great Abaco. It reaches its highest fre-
quency in the interior of the pine forest on
Grand Bahama and in "pure" C. abacoense
from Hole-in-the-Wall on Great Abaco.
Lap-1'' was detected from 4 to 5 localities at
The Crossing where it has a frequency of
about 0.10.

Esterase-2. Woodiuff (1975) first de-
tected variation at this non-specific esterase
locus in C. bendalli from Loc. 243 near

406 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Figure 16. Geographic variation in esterase-2. Sample locality numbers may be discerned from Figure 3.

Snake Cay, Great Abaco. At that time, the
codominant alleles were designated Est-2''
and Est- 2''. Now, as a result of this far more
extensive survey, we report that at least 7
alleles occur at this locus among the Cerion
of the Little Bahama Bank. The various
alleles are all codominant and are desig-
nated Est-^', Est-2'' . . . Est-2'^' in order of
decreasing mobility. The alleles reported
by Woodruff ( 1975 ) are now redesignated
Est-2'' and Est-2^ respectively. Allele fre-
quencies and the overall pattern of allele
distribution are shown in Table 14 and
Fig. 16. Est-2'', a \'ery rare allele found in
only one sample, is omitted from Fig. 16.
The distribution of each allele may now be
considered in turn.

Est-2". A rare allele whose presence is
based on a single specimen of C. bejidalli
from Little Abaco with a two-banded
phenotype interpreted as Est-2VEst-2^

Est-2^. Another rare allele found in 12
samples of C. bendalli from Great and Little
Abaco. Its isolated occurrence on Little
Abaco and near Treasure Cay may be due
to recurrent mutation. Its occurrence at fre-
quencies of up to 0.10 in ten samples from
the area of interaction between C. bendalli
and C. abacoense is probably due to other
forces. We note that nine of these samples
were of C. bendalli or intermediate mor-
photype; only one ( Loc. 257 at The Cross-
ing) was referable to C. abacoense. In this
same area, the Est-2'' allele was not detected
at Locs. 259, 520, 521, 254, and 252 where
snails are judged to be t)'pical C. abacoense.

Est-2''. A rare allele (frequency: 0.01-
O.S) detected in heterozygous form in four
populations of C. ])endaUi from western
Grand Bahama.

Est-2''. A common allele detected in most
populations of Cerion from the Little Ba-

Natural History Cerion VIII: A Revision • Gould and Woodruff 407

hama Bank. It occurs at moderate fre-
quencies (0.02-0.23) throughout most ot
the range of the C. bendaUi morphotype.
The notable feature about the distribution
of this allele is that it reaches higher fre-
quencies in the area of interaction between
the two morphotypes on Great Abaco than
elsewhere. In fact, it is the commonest al-
lele at eight localities in this area. Inter-
sample variation in tliis area is also marked
and is probably too great to be due to
sampling error alone. For example, Est-2''
occurs at the three localities on the eastern
side of The Crossing at frequencies of 0.25-
0.50 while on the western side of The Cross-
ing, only 500 m away, the allele is present
at a frequency of 0.85 in two localities. The
absence of Est-2'i at Sandy Point ( Loc. 251 )
is probably not due to sampling error.

Est-2'' is also present throughout northern
Great Abaco although its presence in some
populations was not detected by Gould et
al. ( 1974 ) . We have subsequently rerun
all the Pongo Cai-pet specimens and now re-
port the occurrence of this allele at low fre-
quency. This correction does not alter any
of the conclusions we reached in that paper
about the systematic status of the Pongo
Carpet morphotype.

Est-2''. This allele was detected on Grand
Bahama where it occurs in all populations
sampled (its absence at Loc. 200 is almost
surely due to sampling error) and is the
commonest allele in the three interior sam-
ples.

Est-2f. This is the commonest allele in
the majority of the Cerion populations on
the Little Bahama Bank. It varies in fre-
quency between 0.76-1.00 in samples of
C. bendalU from Little Abaco and Great
Abaco. It is less common in C. bendaUi
from Grand Bahama where it falls to fre-
quencies as low as 0.32-0.42 in the interior
populations dominated by Est-2'". Lower
frequencies were also noted in the area of
interaction between C. bendaUi and C.
abacoense where Est-2'' was the commonest
allele in 7 of 11 samples. This area and the
adjacent populations of C. abacoense is also

characterized by considerable interpopula-
tion variation in the frequency of Est-2^.
This is particularly marked at The Crossing
where the frequency of Est-2^ increases
from 0.05-0.06 in the mottled western sam-
ples to 0.46-0.75 in the ribby eastern sam-
ples.

Est-2^. A rare allele occurring at low
frequency in five samples of C. bendalli
from Grand Bahama and one sample of in-
termediate morphotype from Great Abaco.

C. Genetic Differentiation of Cerion on
the Little Bahama Bank

The patterns of geographic variation in
the six polymorphic enzymes bear little re-
lation to the distribution of the two taxa
recognized on the basis of shell morphology.
There is not a single case of fixation, or even
near fixation, for alternative alleles in the
two taxa. In only 5 out of the 19 poly-
morphic cases is an allele restricted to one
or the other morphotype: Est-2=\ Est-2*',
Est-2'\ Est-2^ and Mdh-2". With the excep-
tion of Est-2'', which is common in the in-
terior of Grand Bahama, these alleles are
all rare in most or all of the samples in
which they were detected. The o\erall im-
pression emerging from these data is that
C. bendaUi and C. abacoense are very simi-
lar to one another genically. This conclusion
was confirmed by calculating the nor-
malized identity of genes (I of Nei, 1972)
between all 41 samples where N > 11. The
values of I obtained for the 820 pairwise
combinations of samples were in the range
0.9451-0.9999. The average similarity was
0.9849. Values of Nei's ( 1972 ) genetic dis-
tance, D, were accordingly very small and
do not exceed 0.0564.

This overall lack of pronounced genetic
differentiation does not mean that local pat-
terns of variation cannot be discerned. On
the contrary we find sporadic occurrences of
alleles that are unique to one group of pop-
ulations or another. There is also a moder-
ate amount of interpopulation variation that
does not appear to be either obviously cKnal
or closely correlated with simple environ-

408 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

Table 15. Values or mean values for Nei's genetic distance (D) between various samples or
GROUPS of samples (N) OF Ccriou hendalli from Grand Bahama (G.B.) and Cerion from elsewhere
on the Little Bahama Bank. Morphotypes abe B (C. hendalli), Ab (C. abacoense) and I (inter-
medi.^te). Note particularly the across-table low values for comparisons involving Rocky
Point and The Crossing — east, and high values for comparisons involving The Crossing — west.

Morph

Grand

Bahama Sample

Sample(s)

211

204

205

202

201

Loc. 211, G.B.

—

Loc. 204, G.B.

.0082

—

Loc. 205, G.B.

.0066

.0032

—

Loc. 202, G.B.

.0033

.0120

.0129

—

Loc. 201, G.B.

.0051

.0184

.0195

.0018

—

Little Abaco

.0161

.0422

.0329

.0157

.0134

Pongo Carpet

.0170

.0411

.0332

.0142

.0124

Treasure Cay

.0176

.0414

.0336

.0143

.0128

Snake Cay

.0180

.0428

.0350

.0151

.0123

Rocky Point

.0082

.0250

.0233

.0076

.0083

The Crossing—

-east

.0101

.0304

.0285

.0083

.0056

The Crossing—

-west

.0301

.0440

.0417

.0308

.0299

SancK Point

.0170

.0389

.0332

.0141

.0107

Hole-in-the-Wall

.0146

.0356

.0292

.0162

.0154

mental parameters. Regional differentiation
is most marked for the Grand Bahama popu-
lations. These are distinguishable from
Abaconian populations on the basis of their
higher frequencies of Mdh-1" and Lap-1''
and lower frequency of 6-Pgdh-r\ This
differentiation does not, however, permit
the characterization of individual speci-
mens. Only in Est-2 has any regional
diflerentiation of diagnostic genotypes oc-
curred: Est-2'' and Est-2'' are restricted to
Grand Bahama, while Est-2-' and Est-2''
have been detected only on Abaco. A sec-
ond potentially diagnostic allele may be
Got-1'' found in C. aJyacoense and popula-
tions of intermediate morphotype on south-
ern Great Abaco. It is regrettable that we
do not know, at this time, whether Got-f^^
characterizes C. abacoense from Mores
Island or whether it was present in the pre-
siuned extinct C. ahacoeme from Grand
Bahama. Indeed C. ahacoeme cannot be
considered properly characterized until
more samples away from the area of inter-
action with C. hendalli have been analyzed.
We are struck by the similarity between
populations of C. houlalli on Grand Ba-
hama and populations of Cerion from the

area of interaction on Great Abaco. This
pattern emerged repeatedly in the distri-
bution of individual alleles: ' 6-Pgdh-l'',
Lap-1'', and Est-2s (Figures 13, 15, 16). It
is also apparent from a comparison of inter-
population genetic distances (Table 15).
In each set of interpopulation distance com-
parisons, we see that Grand Bahamian pop-
ulations are more similar to Abaconian pop-
ulations from the eastern side of the area of
interaction between C. hendalli and C.
ahacocnse than they are to Abaconian Ce-
rion of either taxon collected away from this
area. This pattern is consistent and not
obscured by the slight regional differentia-
tion on Grand Bahama itself. Note also
that D values between the populations on
either side of The Crossing are greater than
those between various isolated populations
of C. hendalli and greater than those be-
tween the "parental" taxa themselves.

This similaritN' between Cerion from
Grand Bahama and the area of interaction
on Abaco was noted again in the pattern of
variation for individual heterozygosity (H).
Table 16 shows that significantly higher
levels of heterozygosity prevail in these two
areas than elsewhere on the Little Bahama

Natural History Cerion VIII: A Revision • Gould and Woodruff 409

Bank. In this case, however, populations
on both sides of The Crossing are charac-
terized by higher vakies of H.

Although we treat the determinants of
these patterns more fully elsewhere ( Wood-
ruff and Gould, in prep.), we can make
some general comments about their rela-
tionship to ecology and genetics of Cerion.
Cerion populations are very variable in
size. While Woodruff's study demes are
moderate (N = approx. 1,000) in size and
contiguous with adjacent demes, this is not
always the case. Cerion's distribution is
typically patchy, and dramatic declines in
abundance occur over a distance of a few
meters. Stochastic processes are undoubt-
edly important in small, isolated popula-
tions. Gene flow is restricted by the low
vagility of the snails themselves but is
nexertheless demonstrable in nature. Re-
current mutation is probably responsible
for some aspects of genie variation in Little
Bahama Bank Cerion. Finally, selection
may play an important role in regulating the
frequency of certain alleles, either directly
or through its action on coadapted, linked
gene complexes.

It is likely that these various stochastic and
deterministic agents act differentially on
the various populations which differ in size
and degree of isolation or exposure to gene
flow. In this context it may be recalled
that we found moderate amounts cf genie
variability in Cerion: mean number of al-
leles per locus lies in the range 1.65-1.70,
frequency of loci polymorphic per popula-
tion ranged from 0.15-0.30, and the fre-
quencv of heterozygous loci per individual
ranged from 0.054-0. 12S. The occurrence
of more \^ariable populations in some areas
may indicate greater environmental hetero-
geneity or perhaps increased levels of gene
flow between partially differentiated pop-
ulations. Alternatively, lower levels of genie
x-ariation elsewhere may indicate environ-
mental homogeneity and reduced levels of
gene flow. It must be remembered, how-
ex er. that higher (or lower) levels of varia-
bilit\ in different areas (as in the case of

Table 16. Genic heterozygosity (H) per in-
dividual FOR VARIOUS SAMPLES AND GROUPS OF

SAMPLES (N) OF Cerion from the Little Ba-
hama Bank.

Sample(s)

( range )

Grand Bahama

10.43

(9.17-12.27)

Little Abaco

6.25

( 6.22-6.28 )

Pongo Carpet

6.01

( 5.67-6.67 )

Treasure Cay

5.30

(4..52-5.71)

Snake Cav

5.37

(5.00-5.74)

Rockv Point

10.46

(8.95-11.50)

The Crossing—

-east

9.63

(8..30-10.60)

The Crossing—

-west

9.35

(9.03-9.67)

Sandv Point

6.40

—

Hole-in-the-Wall

7.00

Cerion from Grand Bahama and The Cross-
ing on Great Abaco) may have quite dif-
ferent determinants. Thus, while the over-
all pattern of genic variation in these Cerion
may appear relatively simple, we should be
alert for the selective development of
slightly different coadapted gene complexes
in different areas. As in the case of an area
effect in Cepaea nemoralis recently restud-
ied by Johnson (1976), we expect much
synergism between history, environmental
selection, and coadaptation.

Finally, we note that the overall genic
similarity of Cerion on Little Bahama Bank
indicates that these populations were not
founded by dozens of independently de-
ri\'ed hurricane-borne propagules. \Vhile
we cannot exclude the possibility of hurri-
cane transport of alleles from elsewhere, we
cannot properly assess the significance of
such occurrence until we have completed
(jur survey of genic variation elsewhere in
the Bahamas and Cuba. Until this infor-
mation is gathered, we prefer to interpret
the pattern of genic variation as a product
of evolution in sAtu, probably during Pleisto-
cene hypothermals when the Little Bahama
Bank was a single large island. The dif-
ferentiation between Grand Bahama and
Abaco could easily have occurred since the
flooding of the bank, when the various
island populations became isolated from
each other. Using Nei's (1975) crude but
useful method of relating electrophoretic

410 BiiJk'tin Museum of Comparative Zoology, Vol. 148, No. 8

data to time of e^'olutiollarv divergence,
where t = 5 X lO-'D ( and taking 0.0150
for D), we find that the Grand Bahamian
and Abaconian populations diverged about
7,500 years ago if rates of genetic cliange
have been constant. This is very close to
the estimated time of submergence for the
bank.

V. DISCUSSION

The preceding genie analysis has con-
siderable bearing on the taxonomic status of
ribby and mottled moiphotypes. We have
shown that patterns of allozyme variation
bear little relation to distribution of the
shell morphotypes. In fact, these taxa are so
similar to one another that if we had never
seen samples from the area of interaction,
we would probably have concluded that the
two morphotypes are genically identical.
Recall that the highest value of D calcu-
lated among 820 comparisons was only
0.0564 ( 7 = 0.9451 ) . The degree of genie
differentiation found among 47 populations
of Cerion on Little Bahama Bank is well
within the limits found among conspecific
populations of comparable land snails.
Greater interpopulation variation has been
detected among the Helix aspersa inhabit-
ing two adjacent city blocks in Bryan, Texas
(Selander and Kaufman, 1975), among
eight pcjpulations of Theha pisaiia in Israel
(Nevo and Bar, 1976), and among ten
populations of Cepaea nemoralis in North
America (Brussard, 1975). As a generaliza-
tion emerging from a rapidly increasing
number of studies, comparison of local
populations typically produces values of D
in the range 0.001-0.01, while subspecific
comparisons exhibit D — 0.004-0.351, and
specific comparisons yield D — 0.05-2.73
(Nei, 1975). In the willistoni group of
DrosophiUi, for example, average values of
D are: 0.03 between geographic popula-
tions; 0.23 between subspecies and semi-
species; and 0.66 between sibling species
(Ayala, 1975). Clearly, on the basis of
these generalizations, we should synonymize
C. bendalli with C. ahacoeme and treat the

Little Bahama Bank Cerion as a single vari-
able species. We choose not to do this for
several reasons.

First, variation in structural gene prod-
ucts tells us nothing, per se, about the devel-
opment of reproductive isolation. Although
a large number of allelic substitutions usu-
ally precede the completion of reproductive
isolation (typically about 20 per 100 loci
( Ayala, 1975 ) ) , there are many exceptions.
Species pairs characterized by very low
values of D include Drosophila persimiJis
and D. pseudoohscura, 0.05 (Prakash,
1969); Thomoimis bottae and T. umhrinus,
0.009-0.054 (Patton et al., 1972; Patton,
1973); and the semispecies of Drosophila
pauUstorum, 0.025 (Richmond, 1972). At
the other extreme, levels of genie divergence
are similar between various sibling species
of Drosophila, 0.67 (Ayala, 1975); humans
and chimpanzees, 0.62 (King and Wilson,
1975); and local populations of a pocket
gopher Geomii-s hursarius, (Rogers' D =
().65-0.89) ( Penney and Zimmerman, 1976).
Values of D do not by themselves permit us
to make unequivocal taxonomic decisions.

Secondly, the genie and morphometric
surveys, taken together, both indicate that
something notable is going on in the area
of interaction between morphotypes. Al-
though we find no increased variability in
shell form, Cerion from this area are signifi-
cantly more variable genically than sam-
ples collected elsewhere. They are poly-
morphic for alleles not found in either ad-
jacent "parental" population (6-Pgdh-l"
and Est-2'' ) . A similar phenomenon was dis-
covered in the hybrid zone between Mus
musculus muscuhis and M.m. doniesticus in
Denmark (Hunt and Selander, 1973). Pop-
ulations in this area also display higher
levels of P and H, as well as increased inter-
sample variation. This is particularly
marked at The Crossing where between
morphotype gene flow is presumably re-
stricted by an intervening hill. Average
values of D between samples on the east
and west side of the hill are 0.0068 and
0.0003 respectively; D values between sam-

Natltral History Cerion VIII: A Revision • Gould and Woodruff 411

pies on either side of the hill average 0.0235.
\\'e tentatively interpret this situation as
an interaction between two partially dif-
ferentiated taxa possessing slightly different
coadapted gene complexes.

Moreover, on New Providence Island, the
same two morphotypes ( under different
names) interact to yield a "classic" hybrid
zone, with unique phenotypes and greatly
increased morphological variability in the
intermediate samples. In fact, wherever the
two morphotypes interact in the Bahamas
(and they do on several islands), the hybrid
zones are marked and narrow. We have
never failed to find some evidence — either
morphological (as on New Providence) or
genetic (as on Abaco and in partly com-
pleted studies of several zones on Long
Island) — of abrupt change, marked dis-
continuity, or greatly increased variability.
The two morphotypes never blend evenly,
and we take this as a sign that their mixtures
involve two at least moderately discordant
entities. We believe that this discor-
dance deserves some taxonomic recogni-
tion above the subspecific level. The mor-
photypes are not mere geographic variants.
(Simple geographic variants do abound as
well; we designate as such the Pongo Carpet
samples of C. hendaUi because their mor-
phological transitions to normal populations
are smooth and because they share with ad-
jacent samples of normal C. hendalli a
genetic anomaly peculiar to their region —
see Gould et al., 1974.) Structural gene
products, in any case, do not control the
alteration of developmental (allometric)
rates that lie at the core of differences be-
tween morphotypes of Cerion. We shall
haxe to learn how to study the genetics
of eukaryotic regulation before the funda-
mental problems of Cerion are resolved.

Finally, we are now studying a series of
hybrid zones involving Cerion of radically
different morphology elsewhere in the Ba-
hamas. Our preliminary electrophoretic
surveys suggest that some of the most dis-
tinctive morphotypes (recognized as sepa-
rate subgenera) of Cerion have differenti-

ated to a lesser extent than semispecies in
groups like DrosophUa icillistoni. Until we
know more about genie variation in Cerion
as a whole, we will treat the mottled and
ribby morphotypes as semispecies. Until
we know more about them and their inter-
actions (repeated under the guise of many
different species names throughout the Ba-
hamas and Cuba), we will recognize C.
hendaUi and C. ohacoense as taxonomic spe-
cies. In doing so we heed Lewontin's (1974)
closing dictum that "context and interaction
are of the essence."

ACKNOWLEDGMENTS

We thank John Hevelin for spending so
much time measuring all the snails so care-
fully; Margaret Wu, Janet Quensen, Dana
Marek and, especially, Sarah Burgess for
their excellent help in the electrophoresis
laboratory; Tom and Amy Schoener and
William Gillis for sharing their expertise in
Bahamian natural history with us. This
work was supported by N.S.F. Grant BMS-
72-02213-A03. Publication was also par-
tially supported by the Coles Fund.

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NOTE ADDED IN PROOF

Since the above study was completed, a quan-
titati\e system of identif\ing allozymes has been
de\eloped ( Woodruff and Burgess, in prepara-
tion). In future papers, Cerion allozymes will be
characterized by their mobilities ( under specified
conditions) relative to the mobilit>- of analogous
allozymes derived from C. ineaniint. C. incanum
from the Florida Keys is an appropriate standard
as it is genetically invariant throughout most of its
range (Woodruff, D.S., 1978, Evolution and adap-
ti\e radiation of Cerion: a remarkabK- di\erse
group of West Indian land snails. Malacologia
17: 223-239). Allozvmes described here as
6-Pgdh-l", Mdh-1", Mdh-2'>, Got-l'>, and Lap-P
are identical in their mobility to those of C in-
canum and henceforth will be designated with the
superscript 1.00 rather than a letter. For example,
6-Pgdh-l" will now be 6-Pgdh-l' "".

In the course of quantifying the relative mobil-
it\- of various allozxmes, an error was discovered in
the scoring of the Est-2 s>stem as reported above.
While C. ahacoense and C. hendalli share a com-
mon set of Est-2 alleles, the allozyme here re-
ported as Est-2' does not have the same mobility
in its commonest fomi in each species. Est-2', the
commonest allozyme of C ahacoense and the pop-
ulations from the areas of interaction, is now
correctly designated Est-2'"". Est-2' of C. hen-

414 Bulletin Museum of Comparative Zoology, Vol. 148, No. 8

(lalli. on the other hand, is now known to migrate and the calculated interspecific genetic distances

a little fuither. ( This common C. hciidalli allele
is also present, though rare, in C. ahacoense. Thus
it remains true that neither species has a imique
allele. ) Consequently, Table 13 and Fig. 14 are in-
correct with respect to their allozyme frequencies

are slight underestimates. This finding does not
significantly change our overall conclusions. A cor-
rected data set, together with three years' additional
data from the zone of interaction, will be reported
in Woodruff and Gould (in prep.).

APPENDIX: LIST OF LOCALITIES

Specimens described in this paper may be found in the Museum of Comparative Zoology, Harvard Uni-
versity. The authors' collection sites are described below. Grid references are to the Grand Bahaiua and
Abaco ( Bahamas 1 : 25,000 series ) map series prepared by the Directorate of Overseas Surveys. Locali-
ties are arranged geographically from west to east. More precise data are available from the authors.

General Area

near Freeport airport

junction of E. Sunrise Hwy and Shear\vater Dr., Lucaya.

near Blair House on Barbary Beach rd.

site in pine forest, Lucaya Estate.

forest site, North Periiueter Parkway, 1.0 km N. of Queens Hwy.

North Perimeter Parkway

Queens Highway, 0.8 kiu E. of Grand Bahama Hwy junction.

High Rock

North Riding Point — site A.

North Riding Point — site B.

The Gap

Loc.

dRef.

Grand Bah

ama—

-C. hcudaUi

212

7293 29382

211

7324 29351

208

7435 29388

204

7423 29436

207

7589 29488

209

7454 28497

205

7.532 29459

202

7703 29483

199

7834 29.591

200

7831 29.599

201

7871 29548

Little Abaco — C. hendalli

213 TE 2202 2980

214 TE 2286 29792

216 TE 2436 29779

Great Abaco — C. hendalli

217 TE 2437 29780

218 TE 2497 29752

224

2.502

29750

223

2510

29741

222

2511

29740

226

2517

29733

220

2528

29723

219

2531

29721

316

2543

29707

228

2725

29524

229

2716

29529

230

2704

29542

231

2694

295.50

233

2707

29495

240

2866

29404

241

2872

29394

247

2970

29383

246

2927

29273

243

2945

29275

245

2949

29275

244

2953

29274

north coast at Crown Haven.
Wood Cay \illage
Little Abaco end of causeway
Abaco.

between Little Abaco and Great

Great Abaco end of causeway between Little Abaco and Great

Abaco.
Great Abaco Highway, 0.3 km W. Cooperstown ( Pongo Carpet site

9 in Gould et al. 1974).
Cooperstown ( Pongo Carpet site 7 ) .
4.85 km N. of Pongo Carpet ( site 6 )
4.75 km N. of Pongo Carpet (site 5)
3.8 km N. of Pongo Carpet (site 4 )
2.4 km N. of Pongo Carpet ( site 3 )
2.1 km N. of Pongo Carpet (site 2)
Pongo Carpet ( site 1 )
Rock Bluff road, Treasure Cay.
Beach Way, Treasure Cav.
Treasure Cay rd., 1.8 km NW of Loc. 229.
lunction Treasure Cay rd and Great Abaco Hwy.
Great Abaco Hwy. 5.9 km S. of Loc. 231.
Bustick Bight

Great Abaco Hwy, 1.1 km SE. of Loc. 240.
John Cash Point, Marsh Harbour.
Fossil locality exposed in road cut.
Snake Cay rd., W. of causeway to Tuggy Cay.
Tuggy Cay
Snake Cay

Natural History Cerion VIII: A Revision • Gould and Woodruff 415

Loc.

d Ref.

Great Abaco — area of interaction

261

2916

29105

260

2909

29095

501

2907

29095

308

2904

29093

307

2902

29092

306

2900

29092

310

2893

29093

309

2888

29098

305

2893

29088

304

2882

29078

311

2874

29074

249

2845

29059

253

2812

28943

255

2809

28931

251

2594

28783

Great Abaco — C. abacoense

259 TE 2873 29066

520 TE 2856 29050

521 TE 2848 29046
254 TD 2818 28942
257 TD 2816 28933
252 TD 2815 28925
250 TD 2813 28617

APPENDIX [continued]
General Area

between C. bendalli and C. abacoense
Cherokee Sound jetty
Casiiarina Point road junction
0.5 km N. of Rocky Point
0.3 km N. of Rocky Point
Rocky Point

0.3 km S. of Rocky Point
appro.x. 1.0 km W. of Rocky Point
1.6 km W. of Rocky Point
2.9 km N. of Loc. 259
1.6 km N. of Loc. 259

NW. corner of Bahama Pahn Shores estates.
Great Abaco Hwy., 8.5 km S. of Cherokee rd. junct.
Chalk Sound jetty.

Great Abaco Hwy., Crossing Rocks estate junct.
Sandy Point.

Bahama Pahn Shores estate gazebo.

2.2 km S. of Loc. 259.

3.0 km S. of Loc. 259.

Crossing Rocks Bay track: east end.

Crossing Rocks estate beach: north end.

Crossing Rocks estate beach: south end.

Hole-in-the-Wall Ughthouse.

us ISSN 0027-4100

SuUetin OF THE

Museum of

Comparative

Zoology

The American Orb-weaver Genera

Colphepeira, Micrathena and Gasteracantha

North of Mexico [Araneae, Araneidae)

HERBERT W. LEVI

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS. U.S.A.

VOLUME 148, NUMBER 9
8 SEPTEMBER 1978

THE AMERICAN ORB-WEAVER GENERA
COLPHEPEIRA, MICRATHENA AND GASTERACANTHA
NORTH OF MEXICO (ARANEAE, ARANEIDAE)

HERBERT W. LEVP

arranged in loops toward the median side
in the posterior median eyes. In some
(Pachygnatha, and also Linijphia) only the
lateral eyes have the canoe-shaped tapetum
left. Tetragnatha have lost all tapetum.
Some arachnologists consider the absence
of epigynum in Pachijgnatha and Tetra-
gnatha a primitive feature and the two
genera ancestral, related to the haplogyne
spiders. Out-comparison ( all relatives have
a canoe-shaped tapetum, and an epigynum)
would indicate that the loss of both struc-
tures may be secondary. To learn more
about these relationships, the tapetum has
to be examined (Figs. 4, 5, 19, 20, 75, 76).

Already Homann's eye studies indicated
that Colphepeira belongs to the Araneinae,
not close to Meta or Theridiosoma as previ-
ously thought. My study of Colphepeiras
o-enitalia confirms Homann's conclusions. A
new observation, not previously reported,
is that Micrathena species have only rem-
the tapetum into two parts facing each nants of a tapetum in the posterior median
other (Figs. 4, 5, 75, 76). This is char- eyes. Perhaps this is only an adaptation to
acteristic for members of the superfamily Micrathenas diurnal habits or perhaps it
Araneoidea and some related families in- will be of use for figuring out phylogenies.
eluding Agelenidae, Gnaphosidae, Clubion- Also of interest are the relatively large

idae, and Amaurobiidae. Meta and Zij- accessory setae below the tarsal claws of
giella have a large canoe-shaped tapetum, Micrathena. Micrathena may be a good ex-
like Theridiidae, however the tapetum ap- perimental animal for studying the han-
pears reduced in Araneus (and also Col- dUng of silk (a subject about which we
phepeira. Figs. 4, 5) with rows of rhabdomes know little), because its setae against which

silk strands are held by the median claws

^Museum of Comparative Zoology. are larger than those of other genera.

Bull. Mus. Comp. Zool, 148(9): 417-442, September, 1978 417

Abstract. Colphepeira has only one species from
the southeastern United States. There are four
species of Micrathena north of Mexico, three com-
mon ones in eastern North America from New-
England to the tropics, one uncommon from Ari-
;| zona to Guatemala. All Micratheria known, per-
haps 50 species, are tropical American, the three
extending their range north belong each to a
different species group. Even though tropical and
widespread in the eastern states, M. mitrata and
M. gracilis appear absent from southern Florida.
All Micrathena species have only a sliver of the
canoe-shaped tapetum left, the latter a character-
istic of most of the superfamily Araneoidea. The
cosmotropical genus Gasteracantha has only one or
two species in the Americas, G. cancriforinis in the
warmer parts of North America.

INTRODUCTION

Homann ( 1950, 1971 ) reported diversity
in secondary eye structure within the spider
family Araneidae. The tapetum of the
secondary eyes is usually canoe-shaped:
with a crease through the middle dividing

418 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Lcvi 419

Plate 2. Gasteracantha cancriformis (Linnaeus) web, spider removed, dusted with cornstarch. Viscid area
17 cm in diameter. Web was built at 45° angle in porch corner of abandoned Florida house: lower left,
floor; above and far right, walls. Notice tufts of silk on frame lines and on some radii.

Colphepeira contains only one small-
sized species and is related to Singa and
perhaps Mangora. MicratJiena and Gaster-
acantha are tropical spiders; Micrathena,
with more than 50 species, are all tropical
American. Three of these have successfully
extended their range from the tropics to
temperate eastern North America (Map 2).
Each of the three belongs to a different
species group. I believe Gasteracantha and
Micrathena to be specialized Araneidae
which have lost some of their palpal scler-
ites secondarily, and Micrathena do not

attack-wrap prey ( Robinson, personal com-
munication). The lack of attack-wrapping
is probably not primitive but a secondary
loss. Micrathena and Gasteracantha orb-
webs have open hubs (Plates 1, 2). Micra-
thena rests in the center of the web in an
unusual position ( Plate 1 ) and controls
web tension. Unlike other araneid genera
Micrathena have strong fourth legs, used
to hold its position in the web (Plate 1).
Both Micrathena and Gasteracantha are
diurnal spiders.

I would like to thank colleagues for

Plate 1. Micrathena gracilis (Walckenaer) in web, Virginia. The lower photograph is about life-size.
Web dusted with cornstarch. Notice the unusual position of the spider in the open hub and the use of the
fourth leg (in upper photograph).

420 Bulletin Museum of Comparative Zoology, Vol 148, No. 9

making collections available. P. H. Arnaud
and R. X. Schick of the California Academy
of Sciences, D. Bixler, J. A. Beatty, J. Carico,
R. Crabill of the National Musenm of Nat-
ural History, C. Dondale of the Canadian
National Collections, S. I. Frommer of the
University of California Riverside collec-
tions, W. R. Icenogle, B. J. Kaston, H.
Dybas and J. B. Kethley of the Field Mu-
seum of Natural History, T. Kronestedt of
the Natural History Museum, Stockholm,
R. E. Leech, G. Uetz and J. D. Unzicker of
the Illinois Natural History Survey, S. C.
Johnson, W. Peck of the Exline-Peck col-
lection, N. Platnick of the American Mu-
seum of Natural History, W. T. Sedgwick,
W. Shear, W. Star^ga of the Polish Academy
of Science, Warsaw, H. K. Wallace, H. V.
Weems of the Florida State Collection of
Arthropods, F. R. Wanless of the British
Museum, Natural History. The mapping
and typing were done by Susan Hunt. The
research and its publication were made
possible by National Science Foundation
grants BMS 75-05719 and DEB 76-15568.

METHODS

The method of examination, study, and
illustration are those of other studies in this
series on North American orb- weavers.
However in Micrathena and Gasteracantha,
measurement of total length is the length
in midline from the anterior margin of the
carapace to between the posterior abdom-
inal humps or spines.

The tapetum of the secondary eyes was
examined by near-vertical illumination
(with a fiber-light) on the eye. The spider
is kept submerged in alcohol and posi-
tioned on washed sand, which permits odd
positions and very minor changes in angles.
( Because of reflections, sand is a poor
background for most observations; the
background should be black. ) The mag-
nification of the stereoscopic dissecting
microscope is about 50 times for the larger
species, 100 times for the smaller. If the
eye lens has become opaque in preservation
the spider can be cleared in clove oil for

examination. The posterior dorsal eyes were
illustrated with the left eye flat and the
right at an angle, anterior is on top ( Figs.
4, 19, 75). The left lateral eyes were illus-
trated diagrammatically, first the anterior
( left ) flat under the microscope, then the
spider was shifted for the posterior eye
(right) flat under the microscope. The
illustrations produced are thus composites
(Figs. 5, 20, 76). The rows of rhabdomes
can be seen in microscope mounts of the
eyes with a compound microscope.

Following American and British diction-
ary definitions but not arachnological
vocabulary, spines are immovable, rigid,
pointed humps or thorns, as found on the
abdomen of Micrathena and Gasteracantha.
The movable heavy setae covering the in-
tegument are called macrosetae.

Colphepeira Archer

Colphepeira Archer, 1941, Geol. Surv. Alabama,
Mas. Paper, 18: 12. Type species Epeira ca-
tawba Banks by original designation. The name
is feminine.

Diagnosis. Colphepeira differs from many
other araneid genera by the closely spaced
eyes (Figs. 1-3). Unlike Mangora and
Singa, it has a hirsute carapace and ab-
domen with short setae on granules and
tlK> shape of the abdomen is higher than
long with posterior dorsal tubercles (Figs.
1, 6, 7). It differs from Mangora by the
lack of the characteristic long trichobothria
on the third tibia.

Description. The carapace, abdomen,
and legs are covered with scattered short
setae. The setae are cylindrical, distally
tapering to a blunt point, basally with a
narrow neck and sitting in the center of a
disc. The sides of the setae have some
blunt teeth (Fig. 9). The eyes are subequal
to each other in the female, and the an-
terior are larger in the male. The posterior
eye row is recurved. The median eyes are
slightly more than their diameter apart, and
the anterior medians are less than two
diameters from the laterals, the posterior
medians 2.5 diameters from the laterals.

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 421

The thorax has a shallow depression in the
female (Fig. 3) and a transverse, pro-
curved mark in the male. The posterior
part of the head is slighUy swollen. The
height of the clypeus is slightly more than
the diameter of the anterior median eyes
(Fig. 2). The sternum, like the carapace,
is lightly sclerotized and slightly granulate.
The chelicerae are weak, without a basal
boss, and have two teeth on the anterior
margin, two on the posterior and denticles
and one tooth in between (Fig. 8). The
chelicerae have a proximal anterior pro-
jection under the clypeus as in Theridiidae
i^Fig. 8). The proximal articles of the legs,
especially the femora, are also slightly
granulate. The first legs are longest, the
fourth second in length, the third shortest.
The metatarsus and tarsus together are
shorter than the patella and tibia. The
abdomen is higher than long with postero-
dorsal and posterior tubercles (Figs. 1, 6,
7). The lung covers are smooth and, like
those of Meta, lack the transverse grooves
found in species close to Aroneus. The
leaflets of the book-lung in a microscope
mount appear to consist of series of parallel
tracheae attached to each other side by side.

The males are hke females, slightly
smaller and have a more distinct, transverse,
thoracic depression and a slightly higher
clypeus, 1.5 diameters of the anterior me-
dian eyes. The endite has a tooth facing a
tooth on the proximal end of the palpal
femur. The distal margin of the first coxa
has a hook that fits into a groove on the
second femur. The legs are not modified
except that the anterior tibia is slightly
sinuous.

Genitalm. The soft epigynum is covered
with setae (Fig. 9) and has a soft annulate
scape (Figs. 9, 10). The openings appear
on the posterior face (Figs. 10, 11). There
are fertilization ducts. The male palpus
has a soft conductor (C in Figs. 14, 15),
bearing a basal tooth, a median apophysis
(M), which has a hook, and a very large
terminal apophysis (A) which covers con-
ductor and embolus (E) distally. The

Colphepeira catawba ^^

■1

Map 1. Distribution of Colphepeira catawba (Banks),
north of Mexico.

terminal apophysis extends and covers most
of the bulb laterally (Figs. 12-15).

Relatiomhip. A similar large terminal
apophysis (A in Fig. 15) is found in some
species of Singa (Levi, 1972) and Mangora
(Levi, 1975). The resemblance of the
palpus (Fig. 14) to that of Singa hamata
(Clerck) is striking in the shape of the
small median apophysis ( M ) , the soft con-
ductor ( C ) , and the large tenninal apophy-
sis (A). The embolus of Colphepeira is
simpler, and Colphepeira lacks a subter-
minal apophysis. Other similarities to Singa
hamata and Mangora are the lightly scler-
otized epigynum, with a soft broadly at-
tached scape and the closely spaced eyes.
The genitalia also resemble those of Cer-
cidia (except for Cercidias large median
apophysis). All these related genera Col-
phepeira, Singa, Mangora, and Cercidia
have the eyes relatively closely spaced,
unlike those of the larger-sized Aranetis,
Micrathena, Gasteracantha and those of
numerous other araneid genera. CoJpJie-
peira, unlike most araneid genera but like
Mangora, does not have distinct contrasting
ventral abdominal marks.

After he examined the tapetum of the
secondary eyes (Figs. 4, 5) Homann (1950)
first reported that Colphepeira is more

422 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

closely related to Araneus than to Meta or
Thcridiosoma.

Distribution. Only one species of Col-
pJiepeira is known, C. cotaaba, found in
the southeastern United States (Map 1).

Colphepeira catawba (Banks)
Figures 1-15; Map 1

Epcira catawba Banks, 1911, Pioc. Acad. Natur.
Sci. Philadelphia, 63: 450, pi. 34, fig. 4, $.
Female holot\pe from Ash\'ille, Buncombe
County, North Carolina in the Museum of Com-
parative Zoology, lost. There is no old E. B.
Bryant catalog card in the file for this species
as there is for other Banks types.

Aranea catawba-. — Roewer, 1942, Katalog der
Araneae, 1: 859.

Colphepeira catawba: — Archer, 1941, Geol. Surv.
Alabama, Mus. Paper, 18: 13, pi. 1, figs. 3, 4,
pi. 2, figs. 1-3. 1953 Amer. Mus. Novitates, no.
1622: 22, figs. 32-34.

Araiicus catawba: — Bonnet, 1955, Bibliographia
Araneorum, 2: 452.

Description. Female from Arkansas: car-
apace black with paired yellowish white
patches lacking pigment. Sternum spotted
black and yellow-white. Coxae yellow-
white. Legs yellow-white with narrow
black rings. Dorsum of abdomen with
paired streaks (Figs. 1, 7). Venter with
black and white spots and no distinct marks.
The abdomen slightly higher than long,
with four posterodorsal tubercles closely
grouped on each side and a pair of tuber-
cles on the posterior face (Fig. 7). Total
length, 3.5 mm; carapace, 1.2 mm long; 1.0
mm wide. First femur, 1.2 mm; patella and
tibia, 1.4 mm; metatarsus, 1.3 mm; tarsus,
0.5 mm. Second patella and tibia, 1.2 mm;
third, 0.8 mm; fourth, 1.0 mm.

Male from Arkansas with color pattern
less distinct than female. Abdomen shape
like that of female. Total length, 1.6 mm;
carapace, 0.9 mm long, 0.9 mm wide. First
femur, 1.0 mm; patella and tibia, 1.2 mm;
metatarsus, 0.6 mm; tarsus, 0.4 mm. Second

patella and tibia, 1.1 mm; third, 0.6 mm;
fourth, 0.7 mm.

Variation. Some specimens have little
black pigment, others are almost completely
black. Total length of females 2.2 to 3.8 ,
mm; carapace 1.0 to 1.3 mm long, 0.9 to

1.1 mm wide. Total length of males, 1.6 to

2.2 mm; carapace 0.9 to 1.2 mm long, 0.9
mm to 1.0 mm wide.

Diagnosis. This species can only be con-
fused with Dolichognatha species, which
are of similar size and appearance. Doli-
cJwgnatha, a relative of Tetragnutha, has
four small tubercles on the abdomen, and
Colplwpeira has four tubercles postero- il
dorsal on each side and in addition a pair
posteriorly (Figs. 1, 6, 7). DoUcJ^ognatha
species have their chelicerae elongate;
Colphepeira do not (Fig. 2). The Doli-
chognatha epigynum has a depression with
a dark spot on each side; Colphepeira has
a fleshy scape (Figs. 9, 10). The DoU-
chognuthu male palpus appears rather
simple, Tetragnatha-]ike, but with a compli-
cated paracymbium. That of Colphepeira
has a terminal apophysis, median apophysis,
and a simple paracymbium (Figs. 12-15).

Natural History. According to Archer
( 1941 ) ColpJiepeira catawba makes its hori-
zontal orb-web near the ground between
tree roots in thin open second-growth woods
with grassy undergrowth. The web is 7 to
9 cm diameter with about 60 spirals and
a small, poorly defined hub. The egg-sac
hangs with debris, suspended on a hori-
zontal line just above the web. The spider
has its retreat under loose bark and feeds
on small ants.

Distribution. Southeastern United States,
Virginia, southern Florida to Sonora (Map

Records. Virginia. Fairfax Co.: Great
Falls, 9 (N. Banks). Georgia. Troup Co.:
West Point, 7 Sept. 1949, "$ (A. Archer).
Florida. Monroe Co.: 2 mi SE of Mara-

Figures 1-15. Colphepeira catawba (Banks) 1-11. Female. 1. Dorsal view. 2. Eye region and chelicerae.
3. Carapace. 4. Posterior median eyes. 5. Left lateral eyes. 6. Abdomen from side. 7. Abdomen, pos-
terior view. 8. Left chelicera from posterior. 9-11. Epigynum (with enlarged seta). 9. Ventral. 10. Pos-

COLPHEPEIRA, MiCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO • Lcvi 423

I terior. 11. Posterior view, cleared. 12-15. Male, left palpus. 12. Mesal. 13. Ventral. 14. Mesa), expanded.
15. Ventral, expanded.

Scale lines. 0.1 mm, except Figures 1-7, 1.0 mm.

Abbreviations. A. terminal apophysis; C, conductor; E, embolus; H, hematodocha; M, median apophysis;
P, paracymbium; T, tegulum.

424 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

thon, 15 Dec. 1962, jiiv. (W. Ivie). Ala-
bama. Cherokee Co.; May's Gulfe, 11 Aug.
1948, 13 Oct. 1949, 9 9 (A. Archer). Tusca-
loosa Co.: Tuscaloosa, 2 Oct. 1941, 9 (A.
Archer). Mississippi. Forrest Co.: Camp
Shelby, 1945-1946, 9, 6 (A. Ajrcher). Wil-
kinson Co.: Centre ville, Jan.-July 1944, 9
(A.Archer). Arkamus. Carroll Co.: Berry-
ville, Aug. 1938, sununer 1941, Sept. 1944.
9, $ (C.Wilton). Texas. Wilbarger Co.:
4 mi NW of Elliott, 21 Oct. 1964, 6 ( K. W.
Haller). Sonora. Guaymas, on beach, 13
Sept. 1966, 9 (J., W. Ivie), not mapped,
received after completion of paper.

Micrathena Sundevall

Micrathena Sunde\all, 1833, Conspectus Arachni-
dum, London, p. 14. Type species Epeira
chjpeata Walckenaer, the only species listed in
"section one" of the genus. The name is fem-
inine. The synonymy problems of generic names
are discussed by Bonnet, 1957 ( Bibliographia
Araneorum, 2: 2858).

Diagnosis. Micrathena females differ from
those of other genera in ha\'ing a smooth,
shiny carapace with a light rim on each
side (Figs. 18, 31, 45, 59) and in particular,
from Gasteracantha, by having the carapace
longer than wide in the female, at times
with pairs of dimples ( Fig. 31 ) or lateral
spines (in tropical species) unlike that of
any other genera. The female abdomen
is usually longer than wide, trapezoidal, or
square armed with spines, sclerites and a
sclerotized ring around the spinnerets (Figs.
17, 30, 44, 58), while that of Gasteracantha
is usually wider than long. Males lack the
carapace rim and the abdominal spines
and have a smooth, sclerotized abdomen
with a ring around the spinnerets. The male
abdomen is longer than wide, not like that
of Gasteracantha. The median eyes are
never projecting as are those of Gastera-
cantha. The posterior legs of both sexes,
especially the femora, are longer than the
anterior legs or subequal in length, unlike
those of most other araneid genera. The
posterior median eyes have the canoe-
shaped tapetum reduced to a very narrow,
lateral sliver. When viewed through the

lens, it may be hidden by the curvature of
the eyeball (Figs. 19, 32, 46, 60). The
mesal side contains rhabdomes without
tapetum, arranged in rows of a variable
number of loops, few, perhaps 5 to 6 in
number in M. gracilis (Fig. 60), about 8
to 9 loops in the other species (Figs. 19,
32, 46 ) . The narrow lateral tapetum is un-
like that of most species of Araneidae ( Fig.
75). The lateral eyes may be separated
from each other by as much as their di-
ameter; the rhabdomes to the sides of the
tapetum are not arranged in rows.

Description. The carapace is smooth and
shiny in the female and has a unique light
rim on each side (Figs. 18, 31, 45, 59).
Posterior median eyes are 1.2 to 1.5 times
the diameter of anterior medians, laterals
subequal or slightly smaller than anterior
medians. The median eyes are separated by
their diameter to 1.5 diameters. The lat-
erals are several diameters from medians,
but may be up to slightly more than their
diameter from each other (Fig. 47). The
height of the clypeus is equal to or slightly
more than the diameter of the anterior
median eyes (Fig. 57). The chelicerae are
slightly longer than wide, strong with three
to four teeth on the anterior margin, three
to four on the posterior. The legs are
usually not banded. However, sometimes
they are slightly lighter in color than the
carapace and sometimes slightly granulated
(especially the long femora) bearing very
short setae. The abdomen is often brightly
colored, always modified with spines, tuber-
cles, or folds. The spinnerets are sur-
rounded by a sclerotized ring.

The males are smaller than the females
and have the abdomen hghtly sclerotized.
In the males, it is greater in length than in
\\'idth, but lacks the spines and tubercles
of the female. Thus it is quite difficult to
associate with the females of the same
species (Figs. 16, 28, 41, 55). The palpal
femur lacks the proximal tooth, and there
is no facing tooth on the endite. The first
coxae sometimes have a hook, sometimes
not; the hook is absent in the four species

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MEXICO • Levi 425

north of Mexico. The distal articles of the
legs may not be modified and only some-
times have macrosetae; in M. funebris the
first femur has macrosetae on the distal end
(Fig. 16).

Genitalia. The epigynum is usually a
heavily sclerotized knob with openings at
the base of the posterior face ( Figs. 21-23,
34-36, 48-50, 62-&4). There is no annulate
scape. Together with the shape of the
abdomen, the epigynum is a diagnostic
feature but has been slighted by previous
authors.

The palpal patella has one macroseta in
M. funebris; in M. gracilis and M. sagittata
the macroseta is present, but small. The
parac\mbium (P in Fig. 40) differs in
different species (Figs. 25, 38, 52, 66),
unlike other araneid genera but as in Zy-
giella. The bulb lacks a terminal apophysis
but has a transparent flap which arises from
the base of the embolus ( E ) and surrounds
it. It may be homologous with the missing
terminal apophysis (Figs. 27, 40, 54, 68).
The embolus tip ( E ) rests in the conductor
(C). The conductor is sometimes a com-
plex sclerite and at its base another sclerite
may appear, the paramedian apophysis
(PM) (Figs. 40, 68). The median apophy-
sis varies greatly in different species: a
sclerotized, split hook in M. funebris ( M in
Fig. 27), the tip sclerotized in M. mitrata
(Fig. 40), forked in M. sagittata (Fig. 54)
and reduced in M. gracilis (Fig. 68). The
sclerites of the palpus are only lightly
sclerotized, unlike the sclerotized epigy-
num, carapace and spines.

Natural History. Micrathena species are
diurnal and the spiders rest in the open
hub of the orb- web ( Plate 1 ) . The spider
hangs in a characteristic position, controlling
the tension of the web while in the hub. The
spinnerets are up, the dorsal surface of the
abdomen parallel to the ground (Plate 1)
and at an angle to the web plane. The
orb has many radii and spirals. No doubt
the long fourth legs are an adaptation to
the unusual position in the web. Unlike
most araneids, Micratherw species do not

attack-wrap (M. Robinson, personal com-
munication). There is no retreat. All spe-
cies, north of Mexico, mature in fall in the
northern part of their range. Little is known
of egg-sacs and life histories.

Distribution. All species known are
American. There may be as many as 50
or more tropical American species, with
only four extending their range into the
temperate area north of Mexico: three in
the eastern United States, one in the South-
west ( Map 2).

Note. In all species, the genitalia are
quite variable, and thus the species are
difficult to delineate. The four species
north of Mexico, however, are not closely
related to each other and are easily sep-
arated.

Key to Female Micrathena North of Mexico

1. Female abdomen with 5 pairs of conical
tubercles (Figs. 58, 59); eastern United
States to South America gracilis

- Female abdomen with 3 or 2 pairs of
spines or tubercles (Figs. 18, 31, 45) — . 2

2(1) Abdomen with only two pairs of pos-
terior conical tubercles (Figs. 30, 31);
carapace with 3 pairs of dimples ( Fig.
31); eastern United States to South
America (Map 2) _.._ mitrata

- Abdomen with 3 pairs of tubercles or
spines ( Figs. 17, 44, 45 ) and carapace
without paired dimples (Figs. 18, 45) . - 3

3(2) Abdomen much wider behind than ante-
riorly and with anterior, lateral and large
posterior, dorsal spines (Figs. 44, 45);
eastern United States to South America
( Map 2 ) ___ sagittata

- Abdomen as \vide behind as in front with-
out anterior dorsal spines or tubercles, but
with dorsolateral, posterior dorsal and
posterior ventral tubercles (Figs. 17, 18);
Arizona, Baja California to Central
America (Map 2) funebris

Key to Male Micrathena North of Mexico

1. Abdomen wider behind than in front
(Fig. 41); median apophysis of palpus
with two branches ( Figs. 53, M in 54 ) ;
paracymbium a recur\'ed hook pointing
dorsally (Fig. 52); eastern United States
to South America ( Map 2 ) sagittata

- Abdomen as wide behind as in front
(Figs. 16, 28, 55); median apophysis and
paracymbium otherwise (Figs. 26, 39,
67 ) - 2

426 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Map 2. Distribution of Micrathena species north of Mexico.

2(1) Abdomen more than twice as long as
carapace ( Fig. 55 ) ; posterior end of
abdomen longer on \enter than dorsum
( Fig. 56 ) ; eastern United States to South
America (Map 2) gracilis

- Abdomen less than 1.5 times as long as
carapace, not longer on venter ( Figs. 16,
28, 29) . -- 3

3(2) First femur with distal macrosetae ( Fig.
16); paracymbium with large granulate
sculpturing (Fig. 25); median apophysis
a semicircular hook on proximal end of
bulb ( Figs. 26, M in 27 ) ; Arizona, Baja
California to Central America ( Map 2 )
fiinebris

— First femur without distal macrosetae
( Fig. 28 ) ; paracymbium smooth and
small ( Fig. 38 ) ; median apophysis with
distal end pointed and bent i)ack (Figs.
.39, M in 40); eastern United States to
South America ( Map 2 ) ..-. mitrata

Micrathetia funebris (Marx in Banks)
Figures 16-27, IVIap 2

Acrosoma fiinehre Mar.\ in Banks, 1898, Proc. Cali-
fornia Acad. Sci., 3rd ser., 1(7): 249. Female
syntypes from Calmilla Mines and Sierra San
Nicholas in the California Academy of Sciences,
destroyed; and two syntypes from Mazatlan in
the Museum of Comparative Zoology, examined.

Acrosoma macidata Banks, 1900, Canadian Entom.,
32: 100. Female holotype from "Arizona" in the
Museum of Comparati\e Zoology, examined.
NEW SYNONYMY.

Micrathena granulata F. P. -Cambridge, 1904, Bio-
logia Centrali- Americana, Araneidea, 2: 532, pi.
50, fig. 12, S ■ Male holotype from Teapa,
Mexico in the British Museum, Natural History,
examined. Reimoser, 1917, Verb. Zool. Bot. Ges.
Wien, 67: 117. Roewer, 1942, Katalog der
Araneae, 1: 958. Bonnet, 1957, Bibliographia

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 427

Figures 16-27. Micrathena funebris (Marx in Banks). 16. Male. 17-24. Female. 17. Lateral. 18. Dorsal.
19. Posterior median eyes. 20. Left lateral eyes. 21-24. Epigynum. 21. Ventral. 22. Posterior. 23. Lateral.
24. Posterior, cleared. 25-27. Male left palpus. 25. Lateral. 26. Mesal. 27. Submesal, expanded.

Scale lines. 0.1 mm; except Figures 16-18, 1.0 mm.

Abbreviations. C, conductor; E, embolus; H, hematodochia; M, median apophysis; R, radix; S, subtegulum;
T, tegulum; Y, cymbium.

428 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Araneoruni, 2: 2870. Chickering, 1961, Bull.
Mus. Comp. Zool., 125(13): 423, figs. 78-82,
$ . NEW SYNONYMY.

Micrathena funehris: — Reimoser, 1917, Verh. Zool.
Bot. Gesell. Wien, 67: 104. Roevver, 1942,
Katalog der Araneae, 1: 958. Bonnet, 1957,
Bibliographia Araneoruni, 2: 2867. Chickering,
1961, Bull. Mus. Comp. Zool., 125(13): 414,
figs. 55-59, 9.

Micrathena maculata: — Reimoser, 1917, Verh.
Zool. Bot. Gesell. Wien, 67: 10. Roevver, 1942,
Katalog der Araneae, 1: 967. Bonnet, 1957,
Bibliographia Araneoruni, 2: 2871. NEW SYN-
ONYMY.

iSlote. Chickering ( 1961 ) already sus-
pected that the male named M. <iranulatci
belonged to the female M. funehris.

Description. Female holotype of M.
maculata. Carapace brown, sternum biack-
brown. Legs much lighter, yellow-brown,
indistinctly banded darker. Abdomen black
with wliite patches ( Fig. 18 ) . The rim of
the carapace is brown. Carapace with a
circular depression in thorax ( Fig. 18 ) .
Abdomen soft with four fleshy extensions
posteriorly and an anterior pair of humps
on each side (indistinct, if viewed from
above, Figs. 17, 18). Total length 7.0 mm,
carapace 2.3 mm long, 1.6 mm wide. First
femur, 2.0 mm; patella and tibia, 2.2 mm;
metatarsus, 1.5 mm; tarsus, 0.7 mm. Second
patella and tibia, 1.9 mm; third, 0.9 mm.
Fourth femur, 2.4 mm; patella and tibia, 2.2
mm; metatarsus, 1.6 mm; tarsus, 0.7 mm.

Male from Sonora: Carapace glossy
brown; legs brown. Dorsum of abdomen
gray with central white spots and a row of
white .spots along lateral margins (Fig. 16);
sides gray; venter with a plate from pedicel
and surrounding spinnerets lightly sclero-
tized and brownish black. First femur with
strong macrosetae at distal end ( Fig. 16 ) .
Sides of abdomen almost parallel ( Fig. 16 ) .
Total length 4.1 mm, carapace 1.7 mm long,
1.1 mm wide. First femur, 1.6 mm; patella
and tibia, 1.5 mm; metatarsus, 1.0 mm;
tarsus, 0.5 mm. Second patella and tibia,
1.4 mm; third, 0.8 mm. Fourth femur, 1.6
mm; patella and tibia, 1.4 mm; metatarsus,
1.1 mm; tarsus, 0.5 mm.

Variation. The palpus of males from

southern Mexico and Guatemala differs in
having a shorter sclerotized portion of the
median apophysis and a more sclerotized
paramedian apophysis. Females vary in
total length 5.8 to 7.2 mm, carapace 2.4 to

2.6 mm long, 1.5 to 1.7 mm wide. Males
vary in total length 4.0 to 4.1 mm, carapace

1.7 to 2.0 mm long, 1.1 to 1.3 mm wide.
Diag^nosis. This species is similar to M.

mitrata. Females differ in their lack of
the paired dimples on the carapace (Fig.
18), and the openings of the epigynum are
not in a depression (Figs. 22, 24). There
is a pair of tubercles anterodorsal on the
abdomen (Fig. 17), lacking in M. mitrata.
The male palpus differs from M. mitrata in
having a large granulate paracymbium
(Fig. 25) and a split sickle-shaped median
apophysis on the proximal end of the palpal
bulb (Figs. 26, M in 27).

Natural History. Specimens have been
collected sweeping weeds at 975 m eleva-
tion in Sonora. The webs in Arizona were
fairly abundant and were found one to two
feet from the ground, attached to stems of
Johnson grass (Sorghum halepense). They
were found near water, at a 825 m elevation
(J. Beatty, personal communication).

Distrihution. From Baja California and
Arizona to Guatemala (Map 2).

Records (north of Mexico). Arizona.
Pima Co.: Sabino Pond, Santa Catalina
Mts., 825 m el. 26 June, 1960, 9 $ ; 10 July
1962, ? $ (J. Beatty).

Micrathena mitrata (Hentz)
Figures 28-40, Map 2

Epeira mitrata Hentz, 1850, J. Boston Natur. Hist.
Soc, 6: 22, pi. 3, fig. 11, $. Syntypes from
North Carolina and Alabama in the Boston So-
ciety of Natural History, destroyed.

Acrosoma mitrata: — Emerton, 1884, Trans. Con-
necticut Acad. Sci., 6: 327, pi. 38, fig. 9, $.
Emerton, 1902, Common Spiders, p. 189, fig.
438. 9.

Acrosoma reduvianum: — McCook, 1893, American
Spiders, 3: 213, pi. 21, figs. 6, 7, 9, i. Not
PIcctana reduviana Walckenaer, 1841 (= M.
gracilis ) .

Micrathcmi mitrata: — P.P. -Cambridge, 1904, Bio-
logia Centrali- Americana, Araneidea, 2: 538.

CoLPHEPEiRA, MiCRATHENA AND Gasteracantha North OF MEXICO • Levi 429

Figures 28-40. Micrathena mitrata (Hentz). 28-29. Male. 28. Dorsal. 29. Lateral. 30-37. Female. 30.
Lateral. 31. Dorsal. 32. Posterior median eyes. 33. Left lateral eyes. 34-37. Epigynum. 34. Ventral.
35. Posterior. 36. Lateral. 37. Posteriodorsal. 38-40. Male left palpus. 38. Lateral. 39. Mesal. 40.
Submesal, expanded.

Scale lines. 0.1 mm; except Figures 28-31, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; P, paracymbium; PM, paramedian apophy-
sis; R, radix; S, subtegulum; T, tegulum; Y, cymbium.

430 Bulletin Miiscwn of Comparative Zoology, Vol. 148, No. 9

Reimoser, 1917. Verhandl. Zool. Bot. Ges. Wicn,
67: 104. Roewer, 1942, Katalog der Araneae, 1:
966. Kaston, 1948, Connecticut State Geol.
Natur. Hi.st. Surv., 70: 220, figs. 694-695, 9,
S. Bonnet, 1957, Bibliographia Araneoruni, 2:
2872.
Micrathcna reduviancr. — Com.stock, 1912, Spider
Book, p. 517, fig. 563, $. Comstock, 1940,
Spider Book, rev. ed., p. 530, fig. 563, 5 . Not
Plcctana redtiviana Walckenaer, 1841 {= M.
gracilis ) .

Description. Female from Virginia:
Carapace brown with white thoracic rim.
Sternum dark brown. Legs brown, shghtly
hghter than carapace. Dorsum of abdomen
white with black marks (Fig. 31). Sides
white with black marks (Fig. 30). Venter
black. There is a thoracic depression and
three pairs of dimples on each side of thorax
(Fig. 31). Abdomen with four short pos-
terior spines (Figs. 30, 31). Total length
5.0 mm, carapace 1.7 mm long, 1.4 mm
wide. First femur, 1.7 mm; patella and
tibia, 1.7 mm; metatarsus, 1.2 mm; tarsus,
0.5 mm. Second patella and tibia, 1.4 mm;
third, 0.9 mm. Fourth femur, 1.7 mm;
patella and tibia, 1.6 mm; third, 1.1 mm;
fourth 0.4 mm.

Male: carapace brown, posterior median
eyes on black spots. Sternum black. Legs
brown. Dorsum black with paired white
pigment spots. Venter black. Carapace
with three pairs of dimples (Fig. 2S). First
coxa with a veiy small hook. Abdomen
rectangular in dorsal outline (Fig. 28).
Total length 3.5 mm, carapace 1.4 mm long,
1.2 mm wide. First femur, 1.3 mm; patella
and tibia, 1.2 mm; metatarsus, 0.9 mm;
tarsus, 0.5 mm. Second patella, and tibia,
1.0 mm; third, 0.7 mm. Fourth femur, 1.4
mm; patella and tibia, 1.0 mm; metatarsus,
0.8 mm; tarsus, 0.5 mm.

Variation. The dorsal abdominal black
marks are smaller in specimens from Guate-
mala and Panama. Females vary in total
length from 4.7 to 6.0 mm long, carapace
1.7 to 2.2 mm long, 1.4 to 1.9 mm wide.
Males vary in total length from 3.0 to 3.7
mm, carapace 1.5 to 1.8 mm long, 1.1 to
1.2 mm wide. The largest female came
from Mexico.

Diagnosis. Unlike other species north of
Mexico M. mitrata has three pairs of
dimples on the carapace ( Fig. 31 ) . The
female differs from M. funebris in the ab-
sence of the anterodorsal tubercle on the
abdomen (Fig. 30) and the presence of
openings of the epigynum in a depression
(Figs. 34, 35). The male palpus has a
smaller, differently shaped, smooth para-
cymbium (Fig. 38), and a median apophy-
sis folded back on itself, its tip sclerotized
(Figs. 39, M in 40).

Natural History. Micrathena mitrata is
found in deciduous forest, woodland, under
trees, sometimes in shrubs and usually in
the shade.

Distribution. From Maine to Wisconsin
and Kansas, south to Mexico and Panama,
but absent from the Florida peninsula
(Map 2).

Micrathena sagittata (Walckenaer)
Figures 41-54, Map 2

Plcctana sagittata Walckenaer, 1841, Histoire
Naturelle des Insectes, Apteres, 2: 174. The
name was applied to Abbot illustration of
Georgia Spiders, p. 8, fig. 50. Photocopy of the
Abbot manuscript in the Museum of Compara-
tive Zoology, examined.

Epeira spinea Hentz, 1850, J. Boston Soc. Natur.
Hist, 6: 21, pi. 3, fig. 9, 9. Syntypes from
Atlantic states in the Boston Society of Natural
History, destroyed.

Figures 41-54. Micrathena sagittata (Walckenaer). 41-42, Male. 41. Dorsal. 42. Lateral. 43. Subadult male.
44-51. Female. 44. Lateral. 45. Dorsal. 46. Posterior median eyes. 47. Left lateral eyes. 48-51. Epigy-
num. 48. Ventral. 49. Posterior. 50. Lateral. 51. Posterior, cleared. 52-54. Male left palpus. 52. Lateral.
53. Mesal. 54. Mesal, expanded.

Scale lines. 0.1 mm; except Figures 41-45, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulum.

COLPHEPEIBA, MlCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO * Lcvi 431

432 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

AcTOSoma bovimim Thorell, 1859, Oefv. Svensk
Vet. Ak. Forh., 16: 301, 9. Female holotvpe
from Alabama, lost ( not in Natural History
Museum, Stockholm).

Acrosoma spinea: — Emerton, 1884, Trans. Con-
necticut Acad. Sci., 6: 326, pi. 38, figs. 5-8, $,
$ ; 1902, Common Spiders, p. 190, figs. 437,
440-442, 9, $, web.

Acrosoma sagittatum: — McCook, 1893, American
Spiders, 3: 214, pi. 21, figs. 8, 9, $, $.

Micrathena sagittata: — F. P. -Cambridge, 1904,
Biologia Centrali-Americana, Araneidea, 2: 536,
pi. 51, figs. 20, 21, $, i. Comstock, 1912,
Spider Book, p. 514. figs. 189, 558-561, 9 , web.
Reimoser, 1917, Verhandl. Zool. Bot. Gesell.
Wien, 67: 140, pi. 9, fig. 29, 9. Fetrunkevitch,
1930, Trans. Connecticut Acad. Sci., 30: 259,
figs. 111-114, 9, i. Comstock, 1940, Spider
Book, rev. ed., p. 527, figs. 189, 558-561, 9,
web. Roewer, 1942, Katalog der Araneae, 1: 967.
Kaston, 1948, Connecticut Geol. Natur. Hist.
Surv'. 70: 219, figs. 690-693, 2028, 9,5, web.
Bonnet, 1957, Bibliogruphia Araneonmi, 2:
2876.

Micrathena comstocki Archer, 1951, Amer. Mus.
Novitates, no. 1487: 10, figs. 15-17, 9. Female
holotype from Royal Palm State Park [Royal
Palm Area, Everglades National Park], Dade
Count\', Florida in the American Museum of
Natural History, examined. NEW SYNONYMY.

Micrathena sagittata emertoni Archer, 1951, Amer.
Mus. Novitates, 1487: 10, figs. 18, 22, 9.
Female holotype from Norvvell, Plymouth Co.,
Massachusetts, in the American Museum of
Natural History. NEW SYNONYMY.

Description. Female from Virginia:
carapace brown, darker on sides of thorax.
Sternum, legs brown. Dorsum of abdomen
white to yellow with black sclcrotized
disks; black anteriorly above carapace and
posterior spines black (Fig. 45). Sides
black with white patches. Venter black
around spinnerets, with paired white
patches. Abdomen with three pairs of
spines, the pcjsterior ones largest (Figs. 44,
45). Total length from between the pos-
terior .spines 8.0 mm, carapace 3.1 mm
long, 2.5 mm wide. First femur, 3.3 mm;
patella and tibia, 3.0 mm; metatarsus, 1.9
mm; tarsus, 0.9 mm. Second patella and
tibia, 2.8 mm; third, 1.6 mm. Fourth
femur, 3.7 mm; patella and tibia, 3.0 mm;
metatarsus, 2.1 mm; tarsus, 0.9 mm.

Male from Virginia: carapace brown.

Posterior median eyes on black spots. Ster-
num, legs brown. Dorsum of abdomen
lilack, white on lateral margin and posterior
white marks. Sides black, venter black and
brown. Posterior median eyes 1.2 diameters
of anterior medians. Laterals subequal to
anterior median eyes. Abdomen trapezoi-
dal, dorsoventrally flattened (Fig. 41).
Total length 4.7 mm, carapace 1.9 mm long,
1.2 mm wide. First femur, 1.6 mm; patella
and tibia, 1.5 mm; metatarsus, 1.0 mm;
tarsus, 0.6 mm. Second patella and tibia,
1.2 mm; third, 0.8 mm. Fourth femur, 1.8
mm; patella and tibia, 1.4 mm; metatarsus,
1.0 mm; tarsus, 0.6 mm.

Variation. The abdomen of the female
may be white to golden orange in color.
The posterior abdominal spines of speci-
mens from southern Florida are longer than
those from more northern areas. Interest-
ingly some Mexican specimens have minute
posteroventral spines like related tropical
species. Females vary in total length 5.4 to

8.6 mm, carapace 2.9 to 3.5 mm long, 2.2 to

2.7 mm wide. Males vary in total length 4.2
to 5.9 mm, carapace 2.0 to 2.5 mm long, 1.3
to 1.6 mm wide.

Diagnosis. Micrathena sagittata females
are recognized by having three pairs of
spines with the posterodorsals the largest
(Figs. 44, 45), and there are no postero-
ventral spines in specimens north of Mexico
as there are in some related tropical species.
The openings of the epigynum are in de-
pressions on the posterior face of a bulge
(Figs. 4.8-51). The male, unlike other
species of the area, has a trapezoidal ab-
domen, widest posteriorly (Fig. 41) and
a distinct, biforked median apophysis ( Figs.
53, M in 54). The paracymbium, unlike
that of other North American species, is
recurved, pointing back ( Fig. 52 ) with a
spur on its side. Juveniles also have a
triangular abdomen (Fig. 43).

Natural History. This species is found on
shrubs in deciduous forest and woods.

Distribution. From southern New Hamp-
shire to Minnesota, Nebraska, south to
Costa Rica.

COLPHEPEIRA, MiCRATHENA AxXD GaSTERACANTHA NoRTH OF MEXICO • Lcvi 433

Micrathena gracilis (Walckenaer)
Plate 1; Figures 55-68; Map 2

Epcira gracilis Walckenaer, 1805, Tableau des
Aianeides, p. 65. "An unpublished species from
Carolina, communicated by M. Bosc."

Plectana 'gracilis Walckenaer, 1841, Histoire Natur-
elle des Insectes, Apteres, 2: 193. The name
is applied to the Abbot illustration of the Spiders
of Georgia nos. 47, 48. Photocopy of the Mu-
seum of Comparative Zoology, examined. 1
consider this the date of the name.

Pleciana rcduviana Walckenaer, 1841, Histoire
Naturelle des Insectes, Apteres, 2: 201. Name
for Abbot illustration Spiders of Georgia no. 49.
Photocopy in Museum of Comparative Zoology,
examined.

Acrosoma matronale C. L. Koch, 1845, Die Arach-
niden, 11: 68, fig. 887. Female from Mexico,
lost (not in Berlin Museum).

Epeira rugosa Hentz, 1850, J. Boston Natur. Hist.
Soc, 6: 21, pi. 3, fig. 10. Type from southern
states in the Boston Natural History Society,
destroyed.

Acrosoma rugosa: — Emerton, 1884, Trans. Con-
necticut Acad. Sci., 6: 326, pi. 38, fig. 10, 9;
1902, Common Spiders, p. 189, fig. 439, ? .

Acrosoma gracile: — McCook, 1893, American
Spiders, 3: 212, pi. 21, figs. 1-4, 9,6-

Micrathena 77iatronalis: — Simon, 1895, Histoire
Naturelle des Araignees 1: 852, fig. 902, 9.

Micrathena gracilis: — F. P. -Cambridge, 1904, Bi-
ologia Centrali- Americana, Araneidea, 2: 528,
pi. 50, fig. 3, pi. 51, fig. 16, 9, 6. Comstock,
1912, Spider Book, p. 516, fig. 562, 9. Rei-
moser, 1917, Verhandl. Zool. Bot. Ges. Wien, 67:
87, pi. 1, fig. 1, 9 . Comstock, 1940, Spider Book,
rev. ed., p. 529, fig. 562, 9 . Roewer, 1942, Kat-
alog der Araneae, 1: 966. Kaston, 1948, Bull.
Connecticut Geol. Natur. Hist. Surv., 70: 219, pi.
33, figs. 688, 689, 9,6- Bonnet, 1957, Biblio-
graphia Araneorum, 2: 2868. Chickering, 1961,
Bull. Mus. Comp. Zool., 125: 421, figs. 72-77,
9, S.

Micrathena nigrior Chamberlin and Ivie, 1936,
Bull. Univ. Utah, biol. ser. 3(5): 58, figs. 134-
135, 9 . Four female syntypes from Barro Colo-
rado Island, Panama Canal Zone, in the Amer-
ican Museum of Natural History, examined.

Description. Female from Virginia:
carapace brown, darker on sides and middle
of thorax. Sternum maculated white and
brown. Legs brown. Dorsum of abdomen
whitish with dark spots and dark brown
sclerotized spots and dark brown spines
(Fig. 59). Sides brown with white spots
and dark brown sclerotized spots. Thoracic

depression small, round (Fig. 59). Dorsvmi
of abdomen with three pairs of spines and
two pairs of posteriorly directed spines
(Figs. 58, 59). Total length 8.5 mm, cara-
pace 3.0 mm long. 2.2 mm wide. First
femur, 2.3 mm; patella and tibia, 2.2 mm;
metatarsus, 1.4 mm; tarsus, 0.9 mm. Sec-
ond patella and tibia, 2.0 mm; third, 1.4
mm. Fourth femur, 2.7 mm; patella and
tibia, 2.2 mm; metatarsus, 1.5 mm; tarsus,
0.8 mm.

Male: carapace brown, thoracic region
darker. Legs brown. Dorsum of abdomen
whitish, venter blackish. There is a round,
circular thoracic depression (Fig. 55).
Total length 4.8 mm, carapace 1.4 mm long,
0.9 mm wide. First femur, 0.8 mm; patella
and tibia, 0.9 mm; metatarsus, 0.6 mm;
tarsus, 0.4 mm. Second patella and tibia,
0.9 mm; third, 0.6 mm. Fourth femur, 1.2
mm; patella and tibia, 0.8 mm; metatarsus,
0.5 mm; tarsus, 0.4 mm.

Variation. The species is quite variable
in color. Females vary in total length 7.0
to 10.8 mm, carapace 2.6 to 3.7 mm long,
1.7 to 2.5 mm wide. Males vary in total
length 4.2 to 5.1 mm, carapace 1.3 to 1.6
mm long, 0.9 to 1.0 mm wide.

Diagnosis. The female can readily be
recognized by the often gray abdomen with
ten spines ( Figs. 58, 59) and by the laterally
flattened tip of the cone of the epigynum
(Figs. 62-65). The male as well as juvenile
males have an elongate abdomen more than
three times as long as wide (Figs. 55, 56),
the palpus has a round hookshaped para-
cymbium (Fig. 66), a complex conductor
( C ) difficult to make out ( the basal lobe is
probably the paramedian apophysis ) and a
minute median apophysis with a filamen-
tous attachment (Figs. 67, M in 68).

Natural History. Micrathena gracilis is
found in dense woods, the web in shaded
areas, often on bushes. A study of the web
was pubhshed by B. E. Dugdale (1969);
the orb observed had 44 radii and about as
many spirals. The orb had a radius of 17
cm.

Distribution. The species is found from

434 Bulletin Museum of Comparative Zoology, Vol 148, No. 9

eastern Massachusetts, Michigan, Wiscon-
sin, Nebraska, Texas, Sonora, to Panama,
absent from southern Florida (Map 2).

Gasteracantha Sundevall

Gcistcracantha Sundevall, 1833, Conspectus Arach-
nidum, p. 14. Gasteracantha cancrifonnis is
the type species since the only other species
name originalK' included is G. hexacantha, a
synon\ni of G. cancrifonnis. The name is
feminine. Dahl, F. 1914, Mitt. Zool. Mus.
Berlin, 7: 235-301. Benoit, 1962, Ann. Mus.
Royal de I'Afrique Centrale, 8 ser., sci. zool.,
112: 1-70. Emerit, 1974, Faune de Madagas-
car, 38: 1-216.

Vibradctlus Chamberlin, 1925, Bull. Mus. Comp.
Zool., 67: 214. Type species by original des-
ignation and monot>py V. carolinus Chamber-
lin (= Ga.steraca)itJia cancriformis).

Note. Sundevall cited Latreille, 1831 as
author of the name Gasteracantha. How-
ever, Ga.steracanthe Latreille ( 1831 Cours
d'Entomologie, p. 530) is a nomen nudum
since no species are included; it is thus an
invalid name since it lacks an indication
(ICZN, Art. 16, V). Bonnet (1957) also
erroneously cites Latreille ( 1831 ) as author.

The list of synonymy of Gasteracantha is
incomplete; a complete list is found in
Emerit, 1974. There are two excellent re-
visions for the genus available, one of
African species by Benoit and one of Mada-
gascan species by Emerit. Both point out
that Gasteracantha .species are generally
variable and difficult to separate. But this
had been noted previously by Dahl ( 1914 )
in his world-wide study.

Diagnosis. The female carapace is al-
most square in outline (Figs. 71, 72) not
longer than wide nor rebordered on the
sides as that of Micrathena. It is high in
front. Unlike related African genera, there

is only one transverse row of black discs
on the anterior of the abdomen dorsum;
the abdomen has one or two pairs of spines
laterally and one pair posteriorly (Figs.
71, 72). There is a sclerotized, central
bulge on the venter of the abdomen of the
female between genital groove and spin-
nerets, not present in Micrathena (Figs. 71,
73). Unlike Micrathena, Gasteracantha has
a large canoe-shaped tapetum in the pos-
terior median eyes (Fig. 75). Together
with Micrathena, Gasteracantha differs
from other genera in having the spinnerets
on a cone or their base surrounded by a
sclerotized annulus (Fig. 73).

Description. Gasteracantha species are
brightly colored. The sclerotized, square
carapace is high in the head region and has
a deep thoracic groove (Figs. 71, 72). The
eyes are subequal, small, the anterior
median eyes their diameter apart, the pos-
terior medians more than their diameter.
All secondary eyes have a canoe-shaped
tapetum (Figs. 75, 76). The laterals on
each side are far from the medians ( Fig.
74 ) . The height of the clypeus equals the
diameter of the anterior median eyes ( Fig.
74). The heavy chelicerae of G. cancri-
formis have five teeth on the anterior
margin, four on the posterior margin. The
legs are short and thick, and the tarsi are
very short. The female abdomen is a sclero-
tized shield with sclerotized lateral spines
and dorsal sclerotized discs.

Males are minute (Figs. 69, 70, notice
different scale) and less often collected.
The median eye area is slightly projecting
(Figs. 69, 70). The male lacks the tooth
on the proximal end of the palpal femur
and a tooth on the endite present in many
Araneidae. None of the coxae and none of

Figures 55-68. Micrathena gracilis (Walckenaer). 55-56. Male. 55. Dorsal. 56. Lateral. 57-65. Female.
57. Eye region and chelicerae. 58. Lateral. 59. Dorsal. 60. Posterior median eyes. 61. Left lateral eyes.
62-65. Epigynum. 62. Ventral. 63. Posterior. 64. Lateral. 65. Posterior, cleared. 66-68. Male left palpus.
66. Lateral. 67. Mesal. 68. Mesal, expanded.

Scale lines. 0.1 mm; except Figures 55-59, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulum.

COLPHEPEIBA, MiCRATHEXA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 435

436 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

the distal articles of the legs are modified.
Since the abdomen lacks the prominent
spines of the females and also tlie ventral
pnjtuberance, males are difficult to asso-
ciate with matching females in other parts
of the world where there are several species.

Genitalia. The epigynum is a heavily
sclerotized projection with a median lobe
( Figs. 77-80). The openings can be seen on
each side of a septum on the posterior face
(Fig. 79). The internal genitalia (Fig. 81)
are difficult to make out because of heavy
sclerotization.

The palpus is relatively simple. In mesal
view of the contracted palpus three sclerites
are visible: distally the filiform embolus
(Figs. 83, F in 84); in the center a round
sclerite with its distal edge folded and
sculptured, the paramedian apophysis
(PM); and proximally the median apophy-
sis (Figs. 83, M in 84). In the expanded
palpus (Fig. 84) the radix (R) becomes
completely free and transparent hematocha-
like material, probably the conductor (C),
appears behind the embolus (E). The
embolus lacks the parallel lobe (perhaps
the terminal apophysis) of Micrathena. In
the expanded palpus, tlie paramedian
apophysis ( PM ) slips down and behind the
median apopliysis ( M ) as result of pressure
from the soft conductor (C) (Fig. 84).
The Mastophora palpus is similar but lacks
a paramedian apophysis (Levi, in press).
The palpal patella lacks strong setae. The
paramedian apophysis (PM) was called
terminal apophysis by Emerit ( 1968a,
1974). This sclerite is in the same position
and of similar appearance as the para-
median apophysis of Acanthepeira and
other genera (Levi, 1976; in press). The
Acanthepeira paramedian apophysis is
doubtless the same structure as that of the
complex palpus of Eriophora (Levi, 1970)
which was studied by Comstock (1910).
The origin of this structure may perhaps
be seen in the Verrucosa palpus (Levi,
1976, figs. 8, 9) in which it appears to be
the basal end of the conductor. It is also

close to the conductor in Wagneriana and
Wixia (Levi, 1976, figs. 69-71, 98). The
hematodocha-like material (C) behind the
embolus (E) is believed to be the con-
ductor, because of similar structures in sim-
ilar positions in Acanthepeira, Wagneriana,
Wixia, and Scoloderus. Since Gasteracantha
lacks a sclerotized conductor and also
additional lobes on the embolus (including
a terminal apophysis), I believe the palpus
to be simplified secondarily.

Relationship. Gasteracantha is close to
Micrathena and also to Mastophora. The
structure of the palpus, particularly the
lateral (rather than proximal) position of
the tegulum (Figs. 83, 84), the mesal posi-
tion of all sclerites, and the presence of the
paramedian apophysis ( PM ) and conductor
suggest close relationship with Acanthe-
peira, Wagneriana, Wixia, and Scoloderus
( Levi, 1976 ) . Further indication of a highly
specialized araneid are the widely separate
eyes, the square carapace (Figs. 71, 72, 74)
and the modified structure of the abdomen.

Natural History. Gasteracantha biology is
better known than that of many other
araneids (Araneus diadeniatus excepted)
thanks to the beautiful researches of M.
Emerit. His many studies on Madagascan
Gasteracantha versicolor are listed in Emerit
(1974).

Species. Gasteracantha is a cosmotropical
genus. As far as we know, there are only
one or perhaps two species in America,
both known to Linnaeus 200 years ago; G.
tetracantha ( Linnaeus ) in the West Indies
and G. cancriformis (Linnaeus) found
from the southern United States to Argen-
tina.

The literature indicates two species in
the Americas, both originally described by
Linnaeus from Jamaica: Gasteracantha
cancriformis and G. tetracantha. Accord-
ing to the literature, G. tetracantha occurs
from California and Arizona to the Greater
Antilles. The California and Arizona rec-
ords come from specimens of the George
Marx collection, well-known for erroneous

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NoRTH OF MeXICO • Levi 437

Gasteracantha cancriformis

Map 3. North American distribution of Gasteracantha cancriformis (Linnaeus).

records. None were found in recent collec-
tions and the species probably does not
occur in the Southwest. (There is a speci-
men in the N. Banks collection from "Cal."
which probably also originated with Marx. )
Although large collections were available
from Jamaica, only one species, G. cancri-
formis, is found and the Linnaeus record
may also be a locality error. Gasteracantha
tetracantha occurs in Puerto Rico, the
Virgin Islands, and the Bahamas to the
north, exactly those areas where G. cancri-
formis is absent. Many specimens appear
to be intergrades having six spines and
only a few have completely lost the anterior
pair. The intergrades come from the north
and the Bahamas, not from the west. I
hope to obtain more specimens from the
region to determine whether there are one
or two species in the West Indies.

Numerous names have been given to
populations of G. cancriformis but as far as
I know there are never two different pop-
ulations overlapping except perhaps in the
West Indies. The niche of the numerous
African Gasteracantha species seems occu-
pied by species of Micrathena in the Amer-
icas.

Gasteracantha cancriformis (Linnaeus)
Plate 2; Figures 69-84; Map 3

Araiiea cancriformis Linnaeus, 1767, Systema

Naturae, 12 ed., p. 1037. Specimens described

from Jamaica, probably lost.
A. hexacantha Fabricius, 1787, Mantissa Insec-

torum, 1: 344. Name given with one line of

description, but no locality.
Gasteracantha velitaris C. L. Koch, 1838, Die

Arachniden, 4: 33, pi. 269, $. Female from

Brazil.
Plcctana elipsoides Walckenaer, 1841, Histoire

Natiirelle des Insectes, Apteres, 2: 155. Name

given to fig. 118, p. 13 of Abbot, Drawings of

the Insects of Georgia in America, photocopy

examined.
Plcctana quinqueserrata Walckenaer. 1841, Histoire

Naturelle des Insectes, Apteres, 2: 157. Female

from Guyana in Walckenaer's collection, lost.
Plcctana sexserrata Walckenaer, 1841, Histoire

Naturelle des Insectes, Apteres, 2: 157. Female

from Cayenne.
Plcctana atlantica Walckenaer, 1841, Histoire

Naturelle des Insectes, Apteres, 2: 167. Female

from St. Domingo.
Gasteracantha ruhiginosa C. L. Koch, 1845, Die

Arachniden, 11: 55, pi. 878. Female from St.

Domingo, West Indies.
Epeira cancer Hentz, 1850, J. Boston Natur. Hist.

Soc, 6: 23, pi. 3, fig. 13, $. Females from

South Garolina and southern Alabama in the

Boston Natural History Society, destroyed.
Gasteracantlw, insulana Thorell, 1859, Oefv. Svensk

438 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Vet. Akad. Forh., 16: 302. Female from Gala-
pagos Islands in the Natural Histor>- Museum,
Stockholm, examined.

Gasteracantha columhiae Giebel, 1863, Z. Ge-
sammt. Xatiuw., 21: 312. A black individual
from Colombia, lost ( not in Halle ( Saale ) with
the Giebel collection).

Gasteracantha kochii Butler, 1873, Trans. Entomol.
Soc. London, p. 169. New name for G. Iicxa-
cantha: — C. L. Koch, 1838, Arachniden, 4, pi.
117, fig. 268. Female from Para [Belem, Bra-
zil].

Gasteracantha oldendorffi Holmberg, 1876, An.
Agric. Rep. Argentina, 4: 143. Female from
Noter del Rio Guayguiraro, [Entre Rios], Ar-
gentina, destroyed.

Gasteracantha caUida O. P. -Cambridge, 1879, Proc.
Zool. Soc. London, p. 284, pi. 26, fig. 7, 9.
Female holotype from Trinidad, West Indies, in
the Hope Museum, O.vford University, Oxford,
not examined.

Gasteracantha raimondii Taczanowski, 1879, Horae
Soc. Entomol. Rossicae, 15: 106, pi. 1, figs. 25,
26, 5 . Five female syntypes from Lima, Choril-
los and Montana de Nancha, Peru, in the Polish
.'KcademN- of Sciences, examined.

Gasteracantha rainwndii unicolor Taczanowski,
1879, Horae Soc. Entomol. Rossicae, 15: 107.
Two females from Lima, Peru.

Gasteracantlia proboscidea Taczanowski, 1879,
Horae Soc. Entomol. Rossicae, 15: 108, pi. 1,
fig. 27, S ■ Two male syntypes from Lima, Peru
in the Polish Academy of Sciences, examined.

Gasteracantha rufospinosa Marx, 1883, Entomol.
Amer., 2: 25, figs, a-f, 9 , S . Female and male
from Crescent City, Florida, lost ( not in Na-
tional Museum of Natural History).

GasteracantJia maura McCook, 1893, American
Spiders, 3: 210, pi. 13, fig. 12, $. Numerous
syntypes, "young and old from California, par-
ticularly the southern part . . . and from the
islands off the coast," lost ( not in Academy of
Natural Sciences, Philadelphia ) .

Gasteracantha cancrifonnis: — McCook, 1893,
American Spiders, 3: 211, pi. 14, fig. 9, $.
F. P. -Cambridge, 1904, Biologia Centrali-Ameri-
cana, Araneidea, 2: 525, pi. 51, fig. 14, 9.
Petrunkevitch 1930. Trans. Connecticut Acad.
Sci., 30: 249, figs. 103, 9, S. Comstock, 1940,
Spider Book, rev. ed., p. 526, fig. 556, 557, 9,
web. Roewer, 1942, Katalog der Araneae, 1:
949. Bonnet, 1957, Bibliographia Araneorum, 2:
1945.

Gasteracantha elliptica Getaz, 1893, An. Inst. Fis.-
geogr. nac. Costa Rica, 4: 105, 9. Female
specimens from around San Jose, Costa Rica,
depository imknown.

Gasteracantha hiolleyi Banks, 1905, Proc. Entomol.
Soc. Washington, 7 : 20, fig. 3, 9 . Female holo-
type from Cocos Island in the Museum of Com-
parative Zoology, examined.

\'ihradellus carolinus Chamberlin, 1925, Bull. Mus.
Comp. Zool., 67: 214, S- Male holotype from
South Carolina in the Museum of Comparative
Zoology, examined.

Description. Female from Florida: cara-
pace, sternum, legs brownish black. Dor-
sum of abdomen whitish, spines orange-
yellow, muscle scars black. Venter black
with white spots; spines and ventral scler-
otized projection, orange. Total length 7.2
mm, carapace 3.2 mm long, 3.0 mm wide.
First femur, 2.5 mm; patella and tibia, 2.6
mm; metatarsus, 1.4 mm; tarsus, 0.8 mm.
Second patella and tibia, 2.4 mm; third, 1.6
mm; fourth, 2.4 mm.

Male from Florida: carapace brownish
black; sternum black. Legs light blackish
brown. Dorsum of abdomen dark gray with
white spots; venter black with ventral
paired white spots. Total length 2.2 mm,
carapace 1.1 mm long, 0.9 mm wide. First
femvu-, O.S mm; patella and tibia, 0.8 mm;
metatarsus, 0.5 mm; tarsus, 0.3 mm. Second
patella and tibia, 0.7 mm; third, 0.4 mm;
fourth, 0.7 mm.

Variation. There are vast differences in
color and shape ( Fig. 85 ) . Hispaniola and
Jamaican specimens may have an orange
carapace and legs. Almost all Florida
specimens have orange spines. While black
specimens occur occasionally throughout
the range, all specimens from Mona Island
(west of Puerto Rico) were black with light
dorsal patches. Texas specimens were
bright yellow in color (washed out in al-
cohol). The narrowest bodies were found
on Mona Island and Jamaica, the widest

Figures 69-84. Gasteracantha cancriformis (Linnaeus). 69, 70. Male. 69. Dorsal. 70. Lateral. 71-81. Fe-
male. 71. Lateral. 72. Dorsal. 73. Abdomen, ventral. 74. Eye region and chelicerae. 77-81. Epigynum. 77.
Anteroventral. 78. Ventral. 79. Posterior. 80. Lateral. 81. Posterior, cleared. 82-84. Male left palpus. 82.
Lateral. 83. Mesal. 84. Mesal, expanded.

COLPHEPEIRA, MiCRATHENA AND GaSTERACANTHA NORTIf OF MeXICO • Levi 439

Scale lines. 0.1 mm; Figures 69-74, 1.0 mm.

Abbreviations. C, conductor; E, embolus; M, median apophysis; PM, paramedian apophysis; R, radix; T,
tegulum.

440 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

"\g^*«4^l^

Figure 85. Geographic variation of Gasteracantha. Locality data clockwise: Torrey Pines State Park, San
Diego Co., California; Laguna Beach, California; Austin, Texas; College Station, Texas; Hattiesburg, Mississippi;
Houston Co., Alabama; Charleston, South Carolina; Gainesville, Florida; New Providence, Bahamas; Naples,
Florida; Mona Island; La Romana, Dominican Republic; Momance, Haiti; Los Llanos, Cuba; Pinar del Rio,
Cuba; Port Antonio, Jamaica; Mandeville, Jamaica; Barro Colorado Island, Panama Canal Zone; Carmelina,
Honduras; Chichen Itza, Yucatan; Atoyac, Jalisco; Most Southern Palm Grove, Cameron Co., Texas;
Brownsville, Texas; La Paz, Baja California.

Scale line: 5.0 mm.

from Texas to Central America. The short-
est stubbiest spines are found in Mona
Island and California; the longest ones in
Florida and the southeastern states. The
most acute spines are found in Cuban speci-
mens. A characteristic of Hispaniola and
southern Florida specimens is that the sec-
ond pair of spines is larger than the first
pair. Since there are clines of these char-
acters in various directions, it is not easy to
segregate subspecies except for island pop-
ulations.

Unusual variations ai^e the additional
round plates found at times on the dorsum
of the abdomen. The specimen from

Laguna Beach, California illustrated (Fig.
85 ) has an extra assymmetrical plate on the
left anterior. Many specimens throughout
the North American range have the median
posterior plate split into two plates ( Texas,
South Carolina, and Dominican Republic,
Fig. 85).

Total length of females is 5.8 to 8.6 mm,
carapace 2.3 to 3.1 mm wide. Total length
of males 1.9 to 2.7 mm, carapace 0.8 to 1.0
mm wide. Size variation is about the same
throughout the southern states.

Species problems. Archer judging by
museum labels, considered specimens from
the western states to be G. serviUei (Guerin)

CoLPHEPEiRA, MicRATHENA AND Gasteracantha North OF Mexico • Levi 441

and those from the eastern states to be G.
cancriformis; with several subspecies. How-
ever collections from Austin, Texas had
both specimens which Archer considered to
belong to the western species and to the
eastern species. Since there is no overlap
of the two forms, but instead intermediates
are found, I consider all to belong to one
species. The eastern specimens were con-
sidered by Archer (unpublished) to belong
to several different subspecies, but almost
as many were labeled as intermediates (e.g.
from northern Florida), as belonging to
the subspecies.

Diagnosis. Females can be distinguished
from West Indian G. tetracantha (Linnaeus)
by the presence of six spines on the ab-
domen (Figs. 71, 72); G. tetracantha has
only four. Males of G. tetracantha have the
sclerites of the palpus, especially the para-
median apophysis, relatively smaller than
those of G. cancriformis (Figs. 83, 84), al-
though all parts are of the same shape.

Natural History. The striking appear-
ance, conspicuous webs and diurnal habits
make this one of the easily collected tropical
spiders. The web is found between branches
on shrubs and even on buildings (Plate 2).
It is made in the morning and is usually
inclined at an angle, sometimes near vertical
( Plate 2 ) . The outer threads are decorated
with flocculent tufts of silk (Marples, 1969)
and the spider rests in the open hub. Young
Madagascan Gasteracantha versicolor may
have a stabilimentum (Emerit, 1968b).

Adult males have been collected in
Florida from November to February; in
Alabama in August; in Texas in April, June,
July, and October and in California during
July. Adult females can be collected
throughout the year in Florida and Texas.

Muma ( 1971 ) found Gasteracantha can-
criformis webs in central Florida orchards
in trees, between trees and also in mixed
mesophytic woods at a height of less than
1 to more than 6 m. The female's web has
10 to 30 spiral lines, the viscid area span-
ning 30 to 60 cm diameter. The prey caught
are flies, moths, and beetles. The spider

completes the Hfe cycle in a year. Females
mature in late fall or early winter and are
found from October to January. Adult males
first appear in October and November
when females are one-third to one-half
adult size. Adult males hang from a single
strand of silk adjacent to a female's web,
one to three per web. The tiny males ac-
complish sperm induction just prior to
courtship. Egg-sacs are flattened ovate
masses of tangled white, yellow, and green
silk, marked with a longitudinal stripe of
dark green silk. They are found on the
underside of leaves adjacent to the webs.

Distribution. Gasteracantha cancriformis
is found from North Carolina to Southern
California south to northern Argentina
( Map 3 ) . Specimens occasionally get trans-
ported; one female was found among fruit
in Seattle.

LITERATURE CITED

Bexoit, p. L. G. 1962. Monographic des
Araneidae — Gasteracanthinae Africains ( Ara-
neae). Ann. Mus. Roy. de TAfrique Centrale,
Sci. ZooL, ser. 8, no. 112: 1-70.

CoMSTOCK, J. 1910. The palpi of male spiders.
Ann. Entomol. Soc. America, 3: 161-185.

Dahl, F. 1914. Die Gasteracanthen des Berliner
Zoologischen Museums und deren geograph-
ische Verbreitung. Mitt. Zool. Mus. Berlin, 7:
235-301.

Dugdale, B. E. 1969. The weaving of an en-
gineering masterpiece, a spider's orb-web.
Natur. Hist. 78: 36-41.

Emerit, M. 1968a. Le comportement sexuel
d'une araignee Argiopidae malgache a grand
dimorphisme sexuel: Gasteracantha versi-
color Walck. (Araneidae). Bull. Mus. Nat.
d'Hist. Natur., 2 ser., 39: 898-907.

. 1968b. Contribution a I'etude de la bio-
logic et du developpement de I'araignee
tropicale Gasteracantha versicolor (Argiopi-
dae). Bull. Soc. Zool. France, 93: 49-68.

1974. Araneidae Gasteracanthinae.

Faune de Madagascar, no. 38: 209.

Homann, H. 1950. Die Nebenaugen der Ara-
neen. Zool. Jahrb. Abt. Anat. und Ontog.,
72: 289-364.

. 1971. Die Augen der Araneae. Ana-
tomic, Ontogenie und Bedeutung fiir die
Systematik ( Chelicerata, Arachnida). Z.
Morphol. Tiere, 69: 201-272.

442 Bulletin Museum of Comparative Zoology, Vol. 148, No. 9

Levi, H. W. 1970. The Ravilla Group of the
Oib\vea\er Ceniis Eriophora in Xorth .\merica
(Araneae: Araneidae). Psyche, 77: 280-302.

. 1972. The orb-weaver genera Singa

and Hypsosinga in America (Araneae: Aranei-
dae). Pfitjche, 78: 229-256.

. 1975. The American orjj-weaxer genera

Larinia, Cercidia and Mangora north of Mex-
ico (Araneae, Araneidae). Bull. Mus. Comp.
Zooh, 147: 99-1.35.

. 1976. The orb-\\ea\er genera Vcrru-

cosa, Acanthcpcira, Wagneriana, Acacesia,
Wixia, Scolodcnis and Alpaida north of Mex-
ico (Araneae: Araneidae). Bull. Mus. Comp.
ZooL, 147(8): 351-391.

. in press. Orb-webs

orb-weavers. Symp. Zool.

and ph\logeny
Soc. London.

MuMA, M. 1971. Biological and behavioral notes
on Gastcracantha cancrifonnis (Arachnida,
Araneidae). Florida Entomol., 54: 345-351.

M.\RPLEs, B. J- 1969. Observations on decorated
webs. Bull. Brit. Arachnol. Soc, 1 : 13-18.

INDEX

\'alid names are printed in italics. Page numbers
refer to main references, starred page numbers to
illustrations.

atlantica, Plectana, 437

biolleyi, Gasteracantha, 438

bovinum, Acrosoma, 432

callida, Gasteracantha, 438

cancer, Epeira, 437

cancriformis, Aranea, 437

cancrifonnis, Gasteracantha, 437, 439*, 440*

carolinus, Vibradellus, 438

catawba, Aranea, 422

catawba, Araneus, 422

catawba, Colphepeira, 422, 423*

catawba, Epeira, 422

Colphepeira, 420

columbiae, Gasteracantha, 438

comstocki, Micrathena, 4.32

elipsoides, Plectana, 437

elliptica, Gasteracantlia, 438

emertoni, Micrathena sagittata, 432

funebre, Acrosoma, 426

funehris, Micrathena, 426, 427*

Gasteracantha, 434

gracile, Acrosoma, 433

gracilis, Epeira, 433

gracilis, Micrathena, 433, 435*

gracilis, Plectana, 433

granvilata, Micrathena, 426

hexacantha, Aranea, 437

insulana, Gasteracantha, 437

kochii, Gasteracantha, 438

maculata, Acrosoma, 426

maculata, Micrathena, 428

matronale, Acrosoma, 433

matronalis, Micrathena, 433

maura, Gasteracantha, 438

Micrathena. 424

mitrata, Acrosoma, 428

mitrata, Epeira, 428

mitrata, Micrathena, 428, 429*

nigrior, Micrathena, 433

oldendorffi, Gasteracantha, 438

proboscidea, Gasteracantha, 438

quinqueserrata, Plectana, 437

raimondii, Gasteracantha, 438

reduviana, Micrathena, 430

redu\'iana, Plectana, 433

redu\ianum, Acrosoma, 428

rubiginosa, Gasteracantha, 437

rufospinosa, Gasteracantha, 438

rugosa, Acrosoma, 433

rugosa, Epeira, 433

sagittata, Micrathena, 430, 431*

sagittata, Plectana, 430

sagitattum, Acrosoma, 432

sexserrata, Plectana, 437

spinea, Acrosoma, 432

spinea, Epeira, 430

unicolor, Gasteracantha raimondii, 438

\elitaris, Gasteracantha, 437

Vibradellus, 434

(US ISSN 0027-4100)

SulUtin

OF THE

seum

Revision of the Genera and Tropical
American Species of the
Spider Family Uloboridae

BRENT D. OPELL

HARVARD UNIVERSITY

CAMBRIDGE, MASSACHUSETTS, U.S.A.

VOLUME 148, NUMBER 10
27 AUGUST 1979

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bridge, Massachusetts, 02138, U.S.A.

REVISION OF THE GENERA AND TROPICAL AMERICAN
SPECIES OF THE SPIDER FAMILY ULOBORIDAE^

BRENT D. OPELL^

Abstract. The cosmopolitan family Uloboridae
contains 17 genera, four of which are newly de-
scribed. Members of five genera construct orb-
webs, members of three reduced webs derived from
an orb, and members of the remaining genera webs
of unknown form. Recognition of 14 synonyms and
nine new species brings to 28 the number of known
species in the nine uloborid genera found south of
the United States (excluding members of the un-
revised genus Miagrammopes). Scanning electron
microscopy shows spiral capture thread to be joined
to radii by fine cribellate fibrils and allows more
precise identification of male palpal structure, fa-
cilitating cladistic analysis of intrafamilial relation-
ship. This analysis shows the simplest of four tra-
cheal patterns to be derived from more extensive
patterns. Males of three genera have a newly dis-
covered stridulatory apparatus, consisting of a file
on the retrolateral endite surface and two or more
picks on the cymbial tip. Uloborids are character-
ized by having broad, retrolateral patellar notches
and a row of trichobothria on each femur. Presence
of similar trichobothrial rows in several Araneidae
genera indicates close relationship of the two fam-
ilies. Discovery of both haplogyne and entelegyne
uloborids challenges the validity of classifying fam-
ilies solely by the degree of genitalic development.

INTRODUCTION

The spider family Uloboridae is rep-
resented throughout the world, but
achieves its greatest diversity of species
and genera in tropical and subtropical
zones. Muma and Gertsch (1964) revised
the North American members of this fam-

' This study was presented to the Department of
Biology at Harvard University in partial fulfillment
of the requirements for the degree of Doctor of Phi-
losophy.

' Department of Biology, Virginia Polytechnic In-
stitute and State Universitv, Blacksburg, Virginia
24061.

Bull. Mus. Comp.

ily and the present investigation began as
a complementary study of the remaining
American species and genera. However,
it soon became apparent that a clear de-
lineation of American genera required a
reexamination of Lehtinen's (1967) com-
prehensive study of world uloborid gen-
era. Results of this phase of study, while
in essential agreement with most of Leh-
tinen's conclusions, both allowed and de-
manded a more critical look at uloborid
intrafamilial phylogeny. Miagrammopes
is the only Central or South American
genus whose species are not included in
this revision. Comprehensive treatment
of this important and interesting genus
will be presented in a future study.

Olivier described the first uloborid in
1789, placing it {Zosis geniculatiis) into
Linnaeus' genus Aranea. The genus
Uloborus was erected by Latreille in
1806 for U. walckenaerius which, be-
cause of its orb-web, was allied with Ar-
aneidae. Walckenaer described the ge-
nus Uptiotes (Hyptiotes) in 1833 and four
years later changed this genus name to
Mithras, placing it in the family Mithras
along with Scytodes and Lycose. When
O. Pickard-Cambridge first described
Miagrammopes in 1869 he suggested
that it was related to Mithras and Ulob-
orus and that these three genera had
"strong affinity with the family Epeirides
. . . tbut] at present I do not feel suffi-
ciently assured ... to justify the estab-
lishment of a new family of Araneidea,
while, at the same time, I confess myself
quite unable to include them in any fam-

Zool., 148(10): 443-549, August, 1979 443

444 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

il\ hitherto-characterized." During the
same year Thorell recognized Ulobori-
nae as a suhfamilv of Epeiroidae (Ara-
neidae). In 1870, 6. Pickard-Cambridge
estal)lished the family Ulol)()rides for the
genera Hijptiotcs and Ulohoru.s and the
family Miagrammopides for Miagram-
mopes thwaitesii and M. ferdinandi. Si-
mon (1874) combined these two flimilies
to form Uloboridae and in 1892 extended
this family to include the subfamilies
Dinopinae {Dinopis and Menneus), Ae-
butininae {Aehutina), Uloborinae {Uloh-
orus and Sybota), and Miagrammopinae
{Mici^^rammopes and Hijptiotes). Pocock
(1900) and Dahl (1904) removed the Dic-
tynidae (Aebutininae) and Comstock
(1913) the Dinopidae from the family
Uloboridae. However, many workers
(e.g., Gerhard and Kastner, 1937; Kaston,
1948; Bonnet, 1959) place these three
families into the superfamily Dictyno-
idea. A historical review of higher-level
spider classification is presented by Bris-
towe (1938), Bonnet (1959), and Lehtinen
(1967) and need not be repeated here.

Most disagreement about placement of
the Uloboridae centers around the crib-
ellate-ecribellate controversy and a study
of the family might be expected to deal
at length with this problem i.e., can true
spiders be divided into one monophylet-
ic group whose members have a cribel-
liuii and another whose members lack
this structure. However, I agree with
Platnick's (1977) conclusion that the cri-
bellum is a primitive character of all true
spiders and that in some groups it has
been lost or transformed to a colulus. For
this reason attention is devoted to other
characters which may prove more useful
in studies of relationship.

ACKNOWLEDGMENTS

I wish to thank Herbert VV. Levi for
suggesting this study and for his encour-
agement and assistance throughout its
duration. His excellent advice was al-
ways available but never imposed. Pro-

ductivity of field studies in Panama and
Colombia was significantly enhanced by
the biological insights, knowledge, and
hospitality of William Eberhard, Yael Lu-
bin, and Mike Robinson. I hope the out-
come of this investigation will focilitate
their valuable studies of uloborid behav-
ior. Fieldwork was financed by National
Science Foundation Grant for Improving
Doctoral Dissertation Research in the
Field Sciences DEB 77-01571. The
Smithsonian Tropical Research Insti-
tute's Barro Colorado Island and Cali,
Colombia fiicilities were used during
these field studies. Victor M. Patiiio
made it possilile to conduct field studies
at the Jardin Botanico near Mateguadua,
Colombia and the Central Hydroelectica
Anchicaya (Valley de Cauca, Colombia)
provided accommodations and access to
grounds. In the early stages of this study
I benefited from discussions with Ray-
mond Forster and in the late stages Nor-
man Platnick's comments proved ex-
tremely valuable. C. Bradford Calloway
introduced me to techniques used in pre-
paring specimens for the Scanning Elec-
tron Microscope. National Science Foun-
dation Set-Up-Grant BMS-7412494
awarded to the Museum of Comparative
Zoology purchases the SEM (operated by
Edward Seling) used in this study. This
publication is made possible by N.S.F.
grant BMS 75-05719 awarded to Herbert
W. Levi.

Specimens used during this research
were made available by the following
persons and institutions: Anna T. da Cos-
ta, Museu Nacional, Rio de Janeiro;
Charles D. Dondale, Biosystematics Re-
search Institute, Ottawa; William G.
Eberhard; Raymond R. Forster, Otago
Museum, Dunedin; Maria E. Galiano,
Museo Argentina de Ciencias Naturales,
Buenos Aires; M. Grasshoff, Sencken-
berg Museum, Frankfurt; Michel Hubert,
Museum National d'Histoire Naturelle,
Paris; John A. Kochalka; Torbjorn Kro-
nestedt, Naturhistoriska Riksmuseet,
Stockhohn; Herbert W. Levi, Museum of

Revision of Uloboridae • Opell 445

Comparative Zoology, Harvard Univer-
sity; Yael Liil^in; Norman I. Platnick,
American Museum of Natural History;
Gisela Rack, Zoologisches Institut und
Zoologisches Museum, Universitiit Ham-
burg; David C. Rentz, California Acade-
my of Sciences; W. Starega, Polska Aka-
demia Nauk Instytut Zoologiezny,
Warsaw; E. Taylor, Hope Department of
Entomology, Oxford University; Univer-
sity of Vermont Collection; F. R. Wan-
less, British Museum (Natural History);
Howard V. Weems, Jr.; Florida State Col-
lection of Arthropods.

It is impossible to acknowledge the
many individuals responsible for collect-
ing the thousands of specimens which
made this study possible, but their con-
tributions are exemplified by those of
Louis Pefia of Santiago, Chile. His small
collection presented to the Museum of
Comparative Zoology contains one new
species as well as males and females of
two species each known previously from
only a single specimen.

METHODS

Specimens from collections were pre-
pared for study with the A.M.R. 1000
scanning electron microscope (SEM) by
cleaning them in an ultrasonic cleaner or
agitating them in ethanol, dehydrating
them in ethanol, and critical-point-drying
them in carbon dioxide. Prior to exami-
nation, specimens were mounted on dou-
ble-sided Scotch tape affixed to a SEM
stub and then sputter-coated, first with
carbon and then with gold-palladium.
Specially constructed SEM stubs, each
consisting of a 4 mm high, 19 mm diam-
eter, polished aluminum cylinder at-
tached to a 20-mm diameter SEM stub
by silver paint were used to collect web
samples in the field. Double-sided Scotch
tape on the cylinder's 2-mm-wide free
rim served to anchor web samples and
the 15-mm diameter cut-out tape circle
placed on the cylinder's floor provided a
nonconducting surface resulting in a
black background when webs were stud-

ied with the SEM. Web samples were
desiccated and sputter-coated (as de-
scribed above) prior to examination. Hyp-
tiotes male palpi were dissected before
being critical-point-dried and Miagram-
mopes palpi after being critical-point-
dried.

All female genitalia were examined
with both a dissecting microscope and a
differential interference contrast com-
pound microscope. The female genital
region of at least one specimen of each
taxon was studied. Some genitalia were
removed and optically cleared with and
mounted in clove oil. In other cases non-
sclerotized tissues were digested with
hot 10% (weight/volume) potassium hy-
droxide and genitalia were then mounted
(often in situ) in Hoyer's medium. The
genital region of Tangaroa was studied
both after being cleared in clove oil and
as its nonsclerotized tissues were slowly
eroded by 5% sodium hypochlorite. The
latter method permitted study of exposed
regions and the small gas bubbles gen-
erated by this process allowed examina-
tion of the passageway leading from the
central spermatheca.

Specimens were prepared tor tracheal
studies by removing dorsal regions of the
carapace and abdomen and digesting
nonsclerotized tissues in hot 10% potas-
sium hydroxide. Resulting preparations
were first examined with a dissecting
microscope in order to study abdominal
patterns and then with an interference
compound microscope in order to verify
these observations and to study cepha-
lothoracic tracheal patterns. In taxa rep-
resented by few specimens, tracheal pat-
terns were determined by removing
epigyna and carefully separating exposed
muscles to determine the number and
size of underlying tracheae entering the
cephalothorax. Specimens with detached
abdomens can be studied in a similar
manner. In nearly all taxa tracheal pat-
terns of both males and females were ex-
amined and were always found to be sim-
ilar. Therefore, in the few genera

446 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

(Purumitra, Orinomana, and Daramu-
liana) where male tracheal patterns were
not studied (because of lack of speci-
mens), these patterns are most probably
similar to those of females.

Webs to be photographed were first
dusted with cornstarch (Eberhard, 1976b).
Most web measurements were made
from enlarged prints of such photographs
taken at a carefully recorded distance.

NATURAL HISTORY

Uloborids are often referred to as
"hackled-band-orb- weavers," although
webs of only half of this family's 16 gen-
era are known and members of only five
of these genera (Waitkera, Ulohorus, Zo-
sis, Octonoha, and Fhiloponella) con-
struct orb-webs. Polenecia females pro-
duce a vertical, slightly concave web and
hang head down at the webs central at-
tachment to a twin support (Wiehle,
1931). To construct this web the spider
first lays down 25 to 32 radii, adds a sin-
gle temporary spiral loop (which re-
mains), and then deposits cribellar silk
along the radii and framework threads.
Hyptiotes also produces an apparently
reduced vertical web which takes the
form of an orb sector consisting of four
radii across which cribellar capture "spi-
rals" extend. Hyptiotes web construction
and prey capture is discussed by several
authors, including Comstock (1913),
Gertsch (1949), and Nielsen (1932).
Members of the genus Miagrammopes
construct either a single horizontal cap-
ture thread with cribellar silk at its center
or a horizontal resting line with one or
several vertical or diagonal cribellar cap-

ture threads (Akerman, 1932; Lubin et
(iL, 1978).

Uloborid orb-webs are similar in con-
struction to araneid webs. The former
employ a cribellate capture spiral and are
usually horizontal, whereas the latter
have an adhesive capture spiral and are
usually vertical. Eberhard (1969, 1971,
1972) discusses the building, structure,
and ecology of the orb- web of Ulohorus
diversus. Uloborid cribellate silk consists
of two axial (warp or supporting) strands
onto which "puffs" of fine cribellate fi-
brils (Plate 1-F) are combed by the cal-
amistrum (Plate 1-B, C) (Comstock, 1913;
Friedrich and Langer, 1969). To accom-
plish this a spider places the tip of one
fourth tarsus on the dorsal surface of the
opposite fourth tarsus, bringing the cal-
amistrum adjacent and parallel to the cri-
bellum (Plate 2-A). Both legs are rapidly
pumped up and down, combing out fine
cribellar silk (with a diameter of about 30
nm and nodes about every 30 to 40 nm
along its length, Plate 2-B) and placing it
onto the axial threads. As Gerhard and
Kastner (1937) and Gertsch (1949) note,
this spinning behavior is similar to that
of most cribellate spiders. Each pumping
motion is presumably responsible for a
single hackled-band "puff as noted by
Friedrich and Langer (1969) and Langer
and Eberhard (1969). The spider uses
the calamistrum of one leg to spin ap-
proximately half the cribellate strand
spanning adjacent radii and then instan-
taneously switches to the other calamis-
trum with no noticeable interruption
(Eberhard, 1972; personal observations).

The cribellate spiral threads extend di-
rectly across radii, forming a junction that
corresponds to Jackson's (1971) "zero"

Plate 1 . A. Ventral view of Zosis geniculatus female fourth tarsus showing erect macrosetae. B. Prolateral view of fourth
metatarsus of Zosis geniculatus female showing one row of small setae and above it the large setae which form the
calamistrum. C. Ventroprolateral view of Zosis geniculatus female fourth metatarsus and tarsus showing calamistrum
and ventral row of erect macrosetae. D. Cribellum of Zosis geniculatus female. E. Spinning spigots of Fhiloponella
republicana female cribellum. F. Cribellate silk and spiral-radius junction of Uloborus penicillatus female. G. Spiral-
radius junction of Uloborus penicillatus female. H. Spiral-radius junction of Mangaroa sp. female. Horizontal strand is
spiral.

Revision of Uloboridae • Opell 447

448 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

type (Plate 1-F, G). Elierhard( 1976a) has hub's central mesh (Wiehle, 1927; pei-

shown that these junctions, in contrast to sonal observation). This species may also

"nonzero" junctions of araneid orb- add a broad, linear stabilimentum to the

webs, do not allow the spiral to slip when web. As Eberhard (1973) points out, spi-

it is stretched. ral and linear stabilimenta probably

Scanning electron micrographs of serve to conceal the spider from preda-
Ulohorus penicillatus (Plate 1-F, G) tors. However, the flimsy stabilimenta
show the spiral flattened at its junction constructed by Zosis geniculatus seem
with the radius and the cribellate fibrils ill suited for this purpose,
along this junction entangling the radius When the spider senses a prey in its
and responsible for attachment. This is in orb-web it usually jerks the web several
contrast with the spiral of Mangaroa times forcefully (as noted by Marples,
(Plate 1-H) which is attached to the ra- 1962) in a manner reminiscent of mem-
dius by adhesive. It appears that such ad- bers oi Hyptiotes (Comstock, 1913;
hesive junctions allow the spiral to slip Gertsch, 1949) and Miagrammopes (Lu-
through them whereas the entangled bin et al., 1978). This behavior is some-
junctions of uloborids do not permit this times repeated as the spider approaches
to occur. Eberhard (1976a) has shown its prey and may serve to further entangle
that the cribellate mat can pull free of its the prey, evaluate its size, or assure that
rigid axial threads, affording the uloborid it is securely held by the web and will
spiral some elasticity when stressed. This not attack the spider. As the spider nears
movement may allow prey to come into the prey, one of its first legs often probes
contact with additional spiral strands, and lightly contacts the prey by swinging
Perhaps the ability of a spiral to easily in a small circle similar to the exploratory
slide through its radius connections is movements that Eberhard (1972) de-
more important in vertical webs where scribes during spiral construction. Main-
additional spiral strands wait below to in- taining its distance, the spider turns away
tercept the sagging prey. from the prey, hangs from its first and sec-

Stabilimenta are often added to the ond legs and uses its fourth legs to throw
web after the capture spiral has been silk onto the prey. This silk often does
completed and take a variety of forms, not encircle the prey but soon entangles
none of which appear restricted to a sin- its thrashing appendages. After a short
gle genus or species. The most common time the spider moves closer and begins
type of stabilimentum seems to be a lin- a thorough wrapping of the prey which
ear silk band (occasionally two or more) may take from about one minute for a
of variable length which extends through small fruit fly to 20 minutes for an insect
the web's hub, usually with a central gap as large as the spider. This wrapping,
about as long as the spider's body. This conducted while prey is held and manip-
type of stabilimentum is found in webs ulated with the second and third pairs of
of U. diversus (Eberhard, 1969, 1973), LL legs, begins with the struggling prey still
glomosus (Emerton, 1902), V. plumipes in the web's mesh and continues as the
(Wiehle, 1927), PJiilojionella tingena, prey is partly or completely cut or torn
and P. repuhlicana (personal observa- from the web (larger prey are often left
tions). Some U. diversus and U. penicil- suspended by a thread at either end of
latiis add a small spiral of dense silk to the body and rotated on this axis during
the hub of their webs (Eberhard, 1973; wrapping). The spread first legs support
personal observation). Zosis genicuhitus the spider and the fourth legs manipulate
adds either a thin mat of silk to the hub silk used for wrapping. The prey is then
of its web or several delicate, circular silk carried to the web's hub where addition-
cogwheels around the perimeter of the al layers of silk may be added before the

Revision of Uloboridae • Opell 449

^^"^'^Sa^T-

Plate 2. A. Zosis geniculatus female spinning cnbellate silk. B. Cribellate silk strands of Uloborus penicillatus female.
C. Retrolateral view of Philoponella republicana female first patella showing unsclerotized notch (left) bordering lyriform
organs. D. First tarsal organ of Zosis geniculatus female.

spider grasps the prey with its palpal
claws and begins to feed. Members of
this family, lacking poison glands, both
kill and digest prey by pouring digestive
enzymes onto it. The prey's thick silk
wrapping becomes transparent as it ab-
sorbs these enzymes and the extreme
thickness of this covering may be useful
for its absorptive properties rather than
strictly for prey restraint. Uloborids do
not use their chelicerae to pierce or
knead prey and require as much as three
hours to feed on a 5 to 6-mm-long beetle.
Feeding, though slow, is thorough and
when complete, only the prey's exo-
skeleton remains.

Convex eggsacs of Polenecia and Hyp-
tiotes are form-fitted to and nearly indis-
tinguishable from the surfaces of twigs on

which thev are constructed (Comstock,
1913; Scheffer, 1905; Wiehle, 1931).
Members of the genera Uloborus, Zosis,
Philoponella, and Miagrammopes con-
struct suspended, stellate to cylindrical
eggsacs. Philoponella tingena and Zosis
geniculatus first construct a horizontal
silk platform in their webs, deposit a
cluster of 40 to 80 eggs under this plat-
form, and then spin a form-fitting, convex
covering around this egg mass, attaching
its margins to those of the upper platform.
Such eggsacs usually have five to eight
marginal points which represent attach-
ment sites of lines supporting the plat-
form. These observations agree with
those of Uloborus diversus made by
Eberhard (1969). Zosis geniculatus leaves
its thin-walled, pink to light purple egg-

450 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

sac in the orb's mesh where it is seem- appearance of these essentially carefully
ingly ignored. Philoponella tingena fe- constructed webs. Mature males of Phi-
males, on the other hand, remove their loponella, Uloborus, and Zosis also lack
thick-walled, dark gray eggsacs, resus- a cribellum and calamistrum and con-
pend them in a vertical orientation from struct a sheet web similar to that made
the orb's perimeter or framework threads, by second instar spiderlings.
and usually hang immediately above or Since Simon's (1891) description of
beside them (Simon, 1891; pi. 4). It is not Philoponella repuhlicana colonies, little
clear if P. tingena females remain with has been published on this interesting
their eggsacs during the approximately presocial behavior which is apparently
twenty days required for spiderlings to common among members of the genus
emerge, but I observed several females Philoponella. Gertsch (1949) briefly de-
with eggsacs daily for as long as eight scribes this phenomenon, Peters (1953)
days and each time the spider was hang- describes aggregations of P. vicina, and
ing above her eggsac. Uloborus dive rsu.s Struhsaker (1969) notes that P. tingena
females hang their eggsacs at the web's are found in colonies of various sizes and
perimeter (Eberhard, 1969), but U. glo- are sometimes associated with webs of
mosus females arrange their eggsacs in Nephila clavipes. Observations on P. tin-
the web's mesh (Emerton, 1883). Lubin gena show colonies of this species similar
et al. (1978) describe the structure and to those described by Simon,
placement of M/f/^/YZ/nmopt^.S' eggsacs. Philoponella tingena colonies contain
Spiderlings molt once within the egg- as many as 50 individuals of all ages and
sac and emerge as second instars. Newly in larger colonies mature males and fe-
emerged Philoponella tingena spider- males are found in about equal numbers,
lings remained clustered near their egg- A careful search yielded a few solitary in-
sac about a day before ballooning away, dividual s and numerous small colonies of
These spiderlings and a few apparently two of three (usually immature) individ-
third instar spiderlings constructed hori- uals. Colonies were commonly associ-
zontal sheet webs similar to those de- ated with the long, irregular threads of
scribed for Uloborus diversis (Eberhard, Achaearanea webs; however, I found
1969), U. walckenaeriiis and Zosis genie- several individuals and a few small col-
ulatus (Wiehle, 1927), Philoponella vie- onies in irregular scytodid webs and the
ina (Peters, 1953), U. plumipes (Szlep, harrier str-Ands oi Nephila clavipes wehs.
1961), and U. penicillatus (personal ob- Achaearanea webs, often constructed in
servation). Initial stages in construction the protective hollows of buttress tree
of these sheet webs are similar to early roots, appear to provide a ready-made set
stages of orb-web construction seen in of framework threads among which P.
subsequent immature instars and mature tingena can begin building their webs,
females. However, second and a few By removing Achaearanea females to
third instar spiderlings lack a functioning tree trvinks lacking spider webs I was
cribellum and calamistrum and, unable able to study colonization of their newly
to produce a usable capture spiral, begin constructed webs. Not only did second
laying down numerous fine secondary ra- instar P. tingena balloon to these webs
dii after the temporary spiral has been and construct sheet webs, but larger ju-
completed. Spiderlings attach these non- veniles as well as mature males and fe-
sticky radii only at the web's hub and pe- males arrived within six days. One colo-
rimeter. This allows these fine strands to ny of three males, two females, and two
be blown by air currents before they juveniles was established ten days after
come in contact with and attach to pri- the Achaearanea female began con-
mary radii, temporary spiral, and other structing her web.
secondary radii. This explains the sheet Sheet webs of second instar spider-

Revision of Uloboridae • Opell 451

lings and orh-webs of immatiires and ma- tinctive features and suhfamilial affilia-

ture females found with the colony's ii- tion. Based on a study of web structure,

regular framework threads are more or Kaston (1964, 1966) suggests that ulobo-

less horizontal. Mature males do not con- rids evolved along two distinct and pos-

struct webs, but hang from framework sibly polyphyletic lines: one leading via

threads, occasionally taking prey from Sijbota (nomenclatural changes make the

the colony's sheet or orb-webs. Other cited genus Polenecia) to Ulohonis and

than several apparently unsuccessful the other leading via Hyptiotes to Mia-

mating attempts, I observed no close in- grammopes. However, the prevailing

teraction between individuals. Most ju- view of uloborids still seems best ex-

veniles and females hung from the hub pressed by F. Pickard-Cambridge's 1902

of their respective webs and did not clus- statement that: "the spiders compre-

ter in the center of the colony's frame- hended in this family probably have no

work threads as Gertsch describes. Fe- near relations amongst existing forms; so

males with eggsacs hung from framework that it is difficult to place them into any

threads, often those nearest the tree system of classification."

trunk. Whether one views this statement as a

warning or a challenge depends largely

INTRAFAMILIAL PHYLOGENY ^^ ^^^ evaluation and analysis of the fam-
ily's diversity. Three such lines of diver-
Formulation of a clear hypothesis of sity — male and female genitalia and the
uloborid intrafamilial relationship has tracheal system — warrant separate con-
been hindered by the: 1. traditional di- sideration before being used as phylo-
vision of the family into subfamilies Ulo- genetic evidence.

borinae, Hyptiotinae, and Miagrammo- Male Palpus. Despite extensive use
pinae and by emphasizing conspicuous of the uloborid male palpus to distin-
carapace and web modifications of the guish species, no attempt has been made
latter two subfamilies while deemphasiz- to precisely define terms given to its var-
ing less striking, but potentially more sig- ious parts or to assure that these terms
nificant forms of diversity within the Ulo- are used only for homologous structures,
borinae, and 2. failure to carefully and This problem seems to center around use
fully study the family's diversity and em- of the term "conductor" for any sclerite
ploy a clearly defined method for trans- serving to support and orient the embo-
lating this diversity into a phylogeny. lus. By their comparison of the shape and
This is not to say that uloborids have size of the "conductors" of different gen-
been neglected or that those studying era and their failure to specify otherwise,
them have been undiscerning. Rather, Muma and Gertsch (1964), Lehtinen
most studies treat the fauna of a limited (1967), and others imply that these struc-
geographical region and are concerned tures are homologous, when in fact, any
with the immediate task of describing one of five different sclerites may serve
and differentiating species. to support and orient the uloborid em-
By 1966 eight uloborid genera had i^^j^^^ j^ ^^^.^ confusion, I will use the

been described, although, only nve of r .• i . -jc^u £i i

^, . 1 , . 1 T 1 functional term guide for these five anal-

triese were widely recognized. In the ^ ^ , ^ • ^ ^u ^

n . 1 .. J. ^ r^i r ogous structurcs and restrict the term

iirst comprehensive treatment of the fam- ^ , . i • • i

ily since Simon (1892), Lehtinen (1967) conductor to its apparendy original use

described an additional eight genera and ^oi" the specific uloborid sclerite defined

a new subfamily, Tangaroinae. Most of below.

Lehtinen's genera are both valid and In selecting terms to describe the male

monophyletic, but his use of a chart for palpus, I have retained and carefully de-

their description indicates only their dis- fined terms historically associated with

452 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Table 1. Abbreviations used in illustrations.
MALE PALPUS;

STRIDULATORY FILE SF

STRIDULATORY PICK SP

CYMBIAL MACROSETAE CM

FEMORAL TUBERCLE FT

BASAL HEMATODOCHA BH

SUBTEGULUM ST

MIDDLE HEMATODOCHA MH

TEGULUM T

TEGULAR SPUR TS

MEDIAN APOPHYSIS MA

MEDIAN APOPHYSIS BULB MAB

MEDIAN APOPHYSIS SPUR MAS

EMBOLUS E

SPERM RESERVOIR RES

FUNDUS F

CONDUCTOR C

CONDUCTOR BASAL LOBE CBL

CONDUCTOR SPIKE CS

RADIX R

FEMALE GENITALIA;

SPERMATHECA S

FERTILIZATION DUCT FD

ACCESSORY GLAND AG

POSTERIOR PLATE PP

FIGURE SCALE LINES:

a 0.1 mm

b 0.2 mm

c 0.5 mm

d 1,0 mm

Revision of Uloboridae • Opell 453

Plate 3. A. Retrolateral view of Arlston mazolus male left palpus. B. Prolateral view of Ariston mazolus male left palpus.
C. Prolateral view of Siratoba referena male left palpus. D. Apical view of Siratoba referena male left palpus. All scale
lines are 100 fivn long.

the family and introduced additional previously named structures are given

terms only when it was necessary to dis- names which reflect this attachment. Use

tinguish previously unrecognized scler- of terminology common to other spider

ites. Sclerites or projections arising from families does not imply that uloborid pal-

454 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Plate 4. A. Retrolateral view of Hyptlotes gertschi male left palpus. B. Retrolateral view of Hyptiotes gertschi male left
palpus with conductor and median apophysis removed. C. Retrolateral view of Hyptiotes gertschi male left palpus
conductor and median apophysis removed from base of palpus. D. Conductor and median apophysis of Hyptiotes
gertschi male left palpus removed and separated. All scale lines are 100 /xm long.

Revision of Uloboridae • Opcll 455

Plate 5. A. Retrolateral view of Miagrammopes sp. (from Tai Pin) male left palpus. B. Retrolateral view of Miagrammopes
simus male left palpus. C. Apical view of Miagrammopes simus male left palpus with cymbium (attachment at lower
right) removed. D. Apical region of Miagrammopes simus male left palpus separated to show components. All scale
lines are 100 ^m long.

456 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

pal structures are homologous to similar- 1. the free (projecting) embolus (E) con-

ly named structures in these families, taining an ejaculatory duct (ED), 2. a

Rather, the primitive (simple) nature of slender reservoir (RES), and 3. an en-

the palpi of \Vr/;7/c<:'/7/ indicates that most larged fundus (F). Only in Tan^aroa

sclerites of uloborid palpi have evolved (Fig. 19) and possibly in Siratoba and

independently. Ariston, where the embolus is a flat scler-

Endites of Tan^aroa, Waitkera, and ite, is it possible or necessary to distin-

Polenecia have a small stridulatory file guish this sclerite from the ejaculatory

on their lateral surfaces (Figs. 19, 28, 46), duct within. In the remaining genera the

described here for the first time. In Tan- slender, sclerotized embolus closely con-

garoa two and in Waitkera three macro- forms to the ejaculatory duct diameter,

setae, on the distal, ventral cymbial rim Examination of male palpi cleared in

serve as picks (SP) (Figs. 19, 28). Two clove oil did not permit observation of

apparently homologous macrosetae (CM) taenidiae and, therefore, I define the res-

(Plates 6-C, D; 7-C) found on the male ervoir and fundus by size and position

palpi of most other uloborid genera serve rather than by wall structure as does

no obvious function. The palpal femur of Comstock (1910). The embolus arises

Octonoha, Zosis, Philoponelhi, and Ulob- from the dorsal or median tegular surface

orus has one or two proximal, ventral tu- and, when the left tarsus is viewed api-

bercles (FT) (Plate 7-A, C, D). The palpal cally, loops in a clockwise direction

patella and tibia of most genera each (Plates 3-D; 4-B; 5-A, B; 6-B-E, 7-C-D).

have a stout dorsal seta (Fig. 28). In Sy- Radix. A radix (R) arises near the em-

bota and Mia^rammopes a dorsal tibial bolic base and appears to be a primitive

extension protrudes over the tarsus (Plate feature of all uloborids except Tan<iaroa.

5-A, B; Fig. 90). The cymbium of Tan- In Waitkera the radix is a prominent

^aroa folds over the distal part of the tar- scape (Figs. 28, 29) and in Ariston and

sus (Fig. 19), apparently serving as a Siratoba it functions as a guide (Plate 3,

guide for the embolus (E). Figs. 68, 69). The radix of Polenecia, Pu-

Hematodocha and Tefiuhim. The dis- rumitra, Octonoba, and Pacific Philopo-
tal portion of the male palpus of all gen- nelki is a short flange and in the latter three
era bears a basal hematodocha (BH) and genera is usually hidden by the median
sui)tegulum (ST) (Figs. 201, 215, 236). A apophysis (Plate 6-D, E). Remaining gen-
middle hematodocha (MN) and tegulum era lack a radix. Presence of the radix in
(T) is present in Octonoba, Piirumitra, some Philoponelhi and absence in others
Zosis, Philoponelhi, and UloI)orus (Plate indicates that this sclerite's absence in
7-A, D). In the remaining genera the re- this and all other genera but Tangaroa is
gion between the subtegulum and me- the result of loss.

dian apophysis (MA) is not pleated Median Apophysis. In all genera but

(Plates 3-6) and it is unclear whether this Tangaroa a median apophysis arises

region represents a modified subtegu- from the tegulum's center and is usually

lum, hematodocha, or tegulum. For this partially or completely encircled by the

reason it is not named; although, its distal embolus. The median apophysis of Wait-

surface which gives rise to the median /cf'/Y/ consists of a small proximal lobe and

apophysis is termed the tegulum. In Zo- a flat distal portion with a grooved ventral

sis, Octonoba, and Piirumitra a thin teg- surface that forms a guide (Figs. 28, 29).

ular spur (TS) arises from the lateral teg- In Polenecia (Figs. 46, 47) the median

ular surface and serves as an embolus apophysis' distal portion is a flat plate

guide (Plates 6-C; 7-C, D). and the proximal lobe is expanded, serv-

Embolus. In this study I recognize ing along with the conductor (C) as an

three divisions of the male sperm duct: embolus guide. Nearctic members of the

Revision of Uloboridae • Opell 457

Plate 6. A. Retrolateral view of Sybota abdominalis male left palpus. B. Retrolateral view of apex of Sybota osornis
male left palpus. C. Retrolateral view of Octonoba species 1 male left palpus. D, Retrolateral view of Octonoba species
2 male left palpus. E. Apical view of male left palpus of Philoponella sp. from Woodlark Island witfi bulb and conductor
removed to expose embolus and radix. All scale lines are 100 fim long.

458 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

genus Hyptiotes have a long, curved, dis- small, hooked median apophysis spur
tally expanded median apophysis (Plate (MAS), probably homologous with the
4), whereas Palearctic and Oriental mem- distal median apophysis lobe (Plate 7).
bers have a short, flat median apophysis. Conductor. The conductor (C) is a
In all members of this genus the proximal sclerite arising at the junction of the me-
portion of the median apophysis is flat, dian apophysis and tegulum (Plates 3-7).
curved, and tightly appressed to the In several genera the conductor's rugose
grooved conductor to form a guide. This surface aids in its identification (Plates
composite guide is termed the "conduc- 4-C, 6-B). No conductor is present in Tan-
tor" by Muma and Gertsch (1964), Shear oaroa, Purumitra, Octonoba, or Zosis.
(1967), and Lehtinen (1967). Muma and However, in Zosis, Purumitra and Oc-
Gertsch (1964) and Shear (1967) refer to tonoha the small, unsclerotized lobe ex-
the flattened tip of the proximal median tending from the median apophysis
apophysis lobe protruding from the con- bulb's center is probably homologous
ductor groove as the "radix." This term with the conductor (Plates 6-C; 7-C, D).
is unnecessary and inconsistent with the The short, flat conductor of Waitkera
radix as defined above. Members of the (Figs. 28, 29) does not appear to function
genus Siratoba (Plate 3-C, D; Figs. 68, as an embolus guide. Conductors of Po-
69) have a proximal median apophysis lenecia, Hyptiotes, Siratoba and possibly
lobe whose lateral surflice forms a trough Aristori serve in conjunction with the
that accepts the conductor and radix-em- median apophysis' basal lobe as a guide,
bolus unit. In this genus the distal lobe while those of Uloborus, Philoponella,
of the median apophysis is a flat sclerite Ponella, Miagrammopes, and Sybota
with a concave terminus that appears to function independently as a guide. In
accept the tip of the proximal median many Miagrammopes the conductor, in
apophysis lobe (Figs. 68, 69). The medi- addition to forming a trough in which the
an apophysis of Ariston forms an elon- embolus lies, is also modified into a plate
gate projection with an angular terminus of various shapes (Plate 5-B-D). The con-
(Plate 3-A, B). In Sybota the median ductor of Po/enecia (Figs. 46, 47) consists
apophysis is a broad sclerite with two or of a broad basal plate with two distal pro-
three distal projections (Plate 6-A, B). jections, a small dorsal one and a longer,
Miagrammopes is characterized by hav- concave ventral one which, along with
ing an apical median apophysis with two the basal portion of the median apophy-
lobes modified into projections of various sis, serves as a guide. The same condition
shapes which are therefore difficult to if found in Hyptiotes (Plate 4), but here
designate as basal or apical. For this rea- the conductor is thinner and more exten-
son these lobes are designated MA, and sive. In Siratoba (Plate 3-C, D; Figs. 68,
MA2 in Plate 5. Some members of the ge- 69) a Y-shaped conductor (C) lies proxi-
nus Octonoba have a concave median mal to the loop of the radix-embolus unit,
apophysis distal lobe which may serve as apparently supporting it in its fork. The
a guide (Plate 6-C, D). In other members conductor of Ariston (Plate 3-A, B) ex-
the distal region of the median apophysis tends distally under the radix and em-
is rolled, forming a tube (Plate 7-D). Two bolus, but its functional association with
regions can be distinguished in the me- these sclerites is unclear. The conductor
dian apophyses of members of the genera of Miagrammopes, Sybota, and Uloborus
Zosis, Ponella, Philoponella, and Ulobo- (Plates 5; 6-A, B; 7-A) is a trough-shaped
rus: a basal, hemispherical median basal lobe (CBL). In Philoponella, Po-
apophysis bulb (MAB), probably homol- nella, and Miagrammopes a conductor
ogous with the proximal median apoph- spike (CS) is also present (Plates 5-A;
ysis lobe described above, and a distal 7-B).

Revision of Uloboridae • Opell 459

Plate 7. A. Retrolateral view of Uloborus glomosus male left palpus. B. Retrolateral view of Philoponella republicans
male left palpus. C. Retrolateral view oi Zosis geniculatus male left palpus. D. Retrolateral view of Octonoba octonaria
left male palpus with Insert sfiowing tegular spur (small left projection) and embolus. All scale lines are 100 /xm long.

Female Genitalia. The simplest fe- sclerotized mound (Figs. 16, 30). In Tan-
male genitalia are found in Tan^orort and garoa each of the two genital openings
Waitkera where the epigastric furrow's found on the epigastric furrow's anterior
anterior border is formed by a low, un- wall leads to a pair of blind spermathecae

460 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

(S) (Fig. 18). Waitkera females have a
single median spermatheca which opens
only into the vagina through a broad,
hood-shaped dorsal passage (Fig. 31).
Two small, blind, posterolateral accessory
glands open at the posterior margin
of" the genital area. Chamberlain (1946)
describes and illustrates an "epigynal"
opening at each edge of the genital area's
posteromedian indentation, apparently
the openings of the glands described
above.

Members of the genera Polemecia, Ar-
iston, and Hyptiotes have a pair of blind
accessory glands (AG) which appear to
have an external opening distinct from
the spermathecae and which may be
either small (A. albicans and A. mazolu.s)
(Figs. 34, 38) or large (A. ari.statu.s, Po-
lenccia, Hyptiotes) (Figs. 45, 54, 61, 62).
In Ari.ston and Polenecia a pair of blind
spermathecae arises from each genital
opening (Figs. 34, 38, 45, 54). In Hyp-
tiotes a long, coiled duct leads from each
opening and may terminate in a fertiliza-
tion duct (Figs. 61, 62). iMuma and
Gertsch (1964) misinterpret the accessory
glands of Hyptiotes as spermathecae at
the distal end of each long thin coiled
duct. However, VViehle (1927) correctly
recognized the accessory glands and
long sperm ducts as being unconnected.
Females of Aristoti, Polenecia, and
Siratoba have a narrow to broad gen-
ital projection extending over a postero-
ventral concavity in which the openings
are found (Figs. 32, 36, 43, 52, 65, 75). In
Hyptiotes the genital area has a raised
median ventral surface and a broad pos-
terior plate whose narrow, median pro-
jection extends anteroventrally (Figs.
59, 60).

Members of the genera Vloborus, Pu-
rumitra, Zosis, Octonoba, Daramuliana,
Ponella, and Philoponella have similar
internal genitalia, characterized by a pair
of external copulatory openings, each
leading via a duct to a spherical sperma-
theca with a posterior fertilization duct
(FD) (Figs. 139, 161, 180, 190, 200, 210).

Epigyna of Octonoba and Zosis have two
posterior lateral lobes whose posterodor-
sal margins form either a pair of atria
(Figs. 184, 185) or a common atrium
(Figs. 169, 171, 178, 179) in which cop-
ulatory openings are fovmd. A sclerotized
posterior plate (PP) lies dorsal to these
atria (Figs. 171, 185). Uloborus has a sim-
ilar epigynum, but lacks conspicuous
atria, the epigynal copulatory openings
being found at the posterior lobes' pos-
terolateral margins (Figs. 138, 155). Epig-
ynal openings of members of the genera
Daramuliana, Ponella, and Philoponella
lie anterior to the separated or medially
united posterior lobes. In Ponella the
posterior lobes are small mounds (Figs.
198, 199); whereas, in Philoponella and
Daramuliana they appear to form the
posterior rim of an oval or medially di-
vided atrium, respectively, in which cop-
ulatory openings are found (Figs. 189,
208). Members of Puruniitra have two
pairs of sagittally divided median atria
and two pairs of lateral atria (Fig. 160).
Female Sybota, Siratoba, Orinomana,
and Mia^rammopes have a duct (in M.
sinius. Fig. 96, a large bulb and a duct)
leading from each epigynal opening to a
large, blind spermatheca (Figs. 89, 94, 95,
96, 109, 119, 125, 128). At the base of
each spermatheca (or of a thinner duct
leading to it) arises either a fertilization
duct or an expanded region giving rise to
a fertilization duct. Members of Sybota
are charaterized by having an epigynum
with a prominent, well-sclerotized pos-
terior plate, the tip of which is visible in
ventral view as a median extension be-
tween lighter lateral areas at whose an-
terolateral margins the epigynal open-
ings are situated (Figs. 105-108, 112,
113). The epigynum of Orinomana has a
posterior indentation with openings sit-
uated at its lateral margins (Figs. 123,
124, 126, 127). Siratoba has i prominent
epigynal hood which forms a posterior
atrium housing openings at the ventro-
lateral margins of the posterior plate
(Figs. 65, 66, 75, 76). Within the genus

Rfaision of Uloboridae • Opell 461

Miafframmopes s.l. a variety of epigynal
patterns is found, ranging; from a simple
posterior margin with lateral openings
(Fig. 88) to a deeply notched posterior
margin with posterolateral extensions, a
posteromedian scape, and an anterome-
dian rim.

Discovery that members of the genus
Tangaroa, Waitkera, Polenecia, Ariston
and possibly Siratoba and Hijptiotes are
haplogyne {sensus Wiehle, 1967; Cooke,
1969) shows Uloboridae to be an older
lineage than previously thought. Mem-
bers of Tangaroa, Polenecia, and Ariston
lack fertilization ducts and have two pairs
\ of spermathecae, each spermatheca con-
nected by a duct to the genital opening
on the epigastric furrow's anterior wall.
Platnick and Gertsch (1976) and Platnick
(1977) consider two pairs of spermathe-
cae a primitive feature of spiders, but
Platnick (1977) regards such structures in
all non-hypochiloid araneomorphs as a
modified single pair of spermathecae.
Such modification no doubt explains the
multiple spermathecae of many spiders.
However, the striking similarity between
female genitalia of Hi/pochilis (Gertsch,
1958; figs. 14, 16, 20, 21), Hexathele
(Forster and Wilton, 1968; figs. 487-497,
510-514) and those of Tangaroa, Pole-
necia and Ariston (Figs. 18, 34, 38, 45,
54) suggests that genitalia of these three
uloborid genera represent primitive hap-
logyne patterns and are not derived sec-
ondarily.

The single median spermatheca of
Waitkera (Fig. 31) distinguishes it from
other uloborids. However, the male pal-
pal structure of Waitkera (Figs. 28, 29)
suggests that this genus is allied with
Tangaroa and Polenecia and the pres-
ence of a pair of small, presumptive lat-
eral spermathecae indicates that a medi-
an pair of spermathecae may have fused
to form the central spermatheca.

Female genitalia of Oritiomana, Sij-
bota, and most Miagrammopes are simi-
lar in having an anterior pair of blind
spermathecae and a posterior pair of

spermathecae (reduced in Sybota ab-
dominalis) which give rise to fertilization
ducts, making these genera entelegyne.
I hypothesize that these spermathecae
are homologous with those of Tangaroa,
Polenecia, and Ariston and that, with the
possible exceptions of Hijptiotes and Si-
ratoba, fertilization ducts arose only once
in entelegyne uloborids. Implicit in this
hypothesis is homology between the pos-
terior pair of Orinomana, Sybota, and
Miagrammopes spermathecae and the
single pair of spermathecae in the re-
maining entelegyne uloborids.

To my knowledge, no one has attempt-
ed to determine either histologically or
developmentally whether fertilization
ducts represent haplogyne ducts con-
necting the vagina and spermathecae or
if they are newly evolved structures. The
fertilization duct may have evolved in re-
sponse to movement of the haplogyne
genital openings from the deep recesses
of the vagina to the posterior wall of the
epigastric furrow and then to the ventral
surface of the genital region. Only when
these openings moved out of the epigas-
tric furrow and were no longer able to
supply sperm to eggs still within the pro-
tective recesses of the epigastric furrow,
would there be selective pressure for
evolution of a new set of tubes through
which sperm could reach the eggs. This
hypothesis considers female copulatory
openings (into which the male inserts his
embolus) of all spiders homologous and
fertilization ducts apomorphic structures,
derived one or several times.

If, on the other hand, one considers the
fertilization ducts homologous with ducts
connecting the spermathecae and vagina
of haplogyne spiders, one is faced with
the difficulty of explaining how male
mating behavior and palpal structure re-
sponded to a sudden need to use a new
and differently placed aperture for in-
semination. It is difficult to envision a
spider so versatile or a population so di-
verse as to permit this. A similar problem
arises if the fertilization duct is consid-

462 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

ered homologous with one of the paired
lateral spermathecal ducts of haplogyne
spiders and the other duct homologous
with the bursa leading from an external
opening. Only after the two spermathe-
cae were connected by a newly evolved
duct could the external opening be used
by the male. This hypothesis is subject to
the previous criticism and also questions
what selective pressures could be called
upon to explain evolution or movement
of a nonfunctioning aperture.

These conclusions, if correct, support
Shear's (1978) contention that "the hap-
logyne condition represents a grade and
not a clade" by showing that the haplo-
gyne-to-entelegyne transformation has
occurred within a monophyletic lineage.
Belief that Cribellata and Ecribellata rep-
resent separate and monophyletic lin-
eages has been undermined by studies of
Lehtinen (1967), Forster (1968), Baum
(1972), and others and toppled by Plat-
nick's (1977) analysis of this issue. This
being the case, discovery of a single fam-
ily with both haplogyne and entelegyne
members supports Brignoli's (1975) con-
clusions by seriously challenging the va-
lidity of grouping either cribellate or
ecribellate families solely on the degree
of genitalic development.

Tracheal System. Bertkau (1878) re-
ported and Lamy (1902) confirmed that
Vlohorus walckenaerius Latreille had a
stout pair of tracheal trunks which ex-
tended from a common atrium into the
cephalothorax where they divided into
tracheoles which entered the legs. Lamy
(1902) and Petrunkevitch (1933) found a
similar pattern in Mia^,raininopes , but
Petrunkevitch reported that in Ulohorns
^eniculatu.s the two trunks split into
tracheoles upon reaching the petiole. A
study of the Uloboridae shows four basic
tracheal patterns: 1. tracheoles restricted
to abdomen (Fig. 1), 2. tracheoles extend-
ing into cephalothorax (Fig. 2), 3. four
stout tracheal trunks extending into ceph-
alothorax (Figs. 3, 5), and 4. two stout

tracheal trunks extending into cephalo-
thorax (Figs. 7, 9, 11). Within the
cephalothorax only two patterns can be
distinguished: one in which several
small tracheoles serve each leg (Figs. 4,
6, 8, 10) and a second, found only in Mia-
grammopes, in which only one large tra-
chea serves each leg (Fig. 12).

Tracheae always arise from a broad,
common atrium near the cribellum. More
extensive tracheal patterns are not asso-
ciated with forward movement of the spi-
racle as in Hahniidae (Forster, 1970; Pe-
trunkevitch, 1933), some Tetragnathidae
(Levi, 1967), and Anyphaenidae (Plat-
nick, 1974). From the atrium's lateral
margin small tracheae, shown by Purcell
(1909, 1910) to be homologous with book
lungs, extend posteriorly, presumably to
serve silkglands and spinnerets. From
the atrium's anterior margin two stout
median tracheal trunks extend into the
abdomen (pattern 1) or the abdomen and
cephalothorax (patterns 2-4). In the latter
case each median trachea gives rise to a
large lateral branch which divides into
tracheoles that supply the abdomen. In
pattern 3 these lateral branches arise just
posterior to the median trunks' bifurca-
tion.

The generally accepted view that in
spiders an extensive tracheal system
evolved primarily to reduce respiratory
water loss is presented by Levi (1967)
and is well documented for small spi-
ders. Anderson (1970) accepts this expla-
nation, but suggests and provides sup-
porting evidence for the hypothesis that
tracheal systems also evolved to supply
increasing oxygen demands associated
with increasing metabolic rates. Higher
metabolic rates, he suggests, resulted
when more reliable food supplies initi-
ated more active forms of prey capture,
increased fertility and more rapid devel-
opment rates. Citing his findings and
those of Davis and Edney (1952) and
Dresco-Derouet (1960), who discovered
that blockage of the tracheal spiracle
caused no significant decrease in basal

Revision of Uloboridae • Opell 463

(0 o -r-

Q. Q (U

o a: —

464 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

(resting) metabolic rate, Anderson further
hypothesized that it is largely during peri-
ods of activity that tracheae serve to pre-
vent respiratorv' water loss and to provide
large amounts of oxygen. Levi and Kirber
(1976) cite studies supporting the hy-
pothesis that tracheae are particularly im-
portant in supplying oxygen to the ceph-
alothorax during periods of activity
when, in order to generate hydrostatic
pressure for leg extension, hemolymph
circulation between cephalothorax and
abdomen is thought to be stopped. A
comprehensive reivew of studies relating
to hydrostatic pressure and locomotion is
presented by Anderson and Prestwich
(1975) who also present evidence show-
ing that during extreme activity retro-
grade flow and residting accumulation of
hemolymph in the abdomen is attenuat-
ed by: 1. closure of a valve between the
heart and anterior aorta, 2. increased ab-
dominal hemolymph pressure, 3. rigidity
of the book lungs, and 4. large pressure
drops as venous hemolymph passes
through the book lungs. These findings
indicate that circulation of oxygen-car-
rying hemolymph between abdomen
and cephalothorax may not be complete-
ly blocked during activity or, if so, is
stopped for only short intei-vals. Owing
to proportional hemolymph distribution
and maintained abdominal hemolymph
pressure, flow is quickly restored when
activity ceases. Anderson and Prestwich
hypothesize that reliance on cephalotho-
rax-generated fluid pressure for leg ex-
tension allowed more leg muscle mass to
be devoted to flexion, thereby increasing
strength for prey capture. Canying this
reasoning a step further, I suggest that
some advanced tracheal systems may
have evolved largely to supply the in-
creased oxygen demands of these flexor
muscles.

To summarize, an advanced tracheal
system which extends into the cephalo-
thorax may serve to: 1. reduce respiratory
water loss, 2. increase total oxygen sup-
ply, 3. supplement or replace hemo-

lymph-carried oxygen during periods of
activity, and 4. supply oxygen specifical-
ly to leg flexor muscles. Although these
four functions are not mutually exclusive,
it is likely that tracheal patterns found in
different families or even those found
within a single family evolved under se-
lective pressures favoring a certain func-
tion or set of functions.

Habitats, behavior and webs of many
uloborid genera are unknown, but evi-
dence suggests that tracheal patterns 3
and 4 of some genera serve largely to sup-
ply oxygen for extended or particularly
strenuous leg flexure. Members of the
genera Polenecia, Hyptiotes, Miagram-
mopes, construct reduced, highly spe-
cialized webs (see Natural History) and
have tracheal pattern 4. Hyptiotes and
Miagrammopes (perhaps Polenecia as
well) support and flex the web while
waiting for prey and vigorously jerk the
web to ensure prey entrapment. Tracheal
patterns of SyJ)ota and Orinomana,
shown by this study to be closely related
to Miagrammopes, may facilitate similar
specializations of web and behavior.

Among genera known to construct orb-
webs (Waitkera, Zo.si.s, Octonoba, Philo-
ponella, and Ulohorus) all four tracheal
patterns are represented. Size ranges of
these genera are similar with smaller
species belonging to Philoponella (tra-
cheal pattern 1). In Panama and Colom-
bia I collected specimens of Philoponel-
la, Ulohorus (tracheal patterns 3 and 4),
and, less often, Mia gram mopes, in the
same habitat. However, in captivity
members of the latter genus appeared
least resistant to desiccation and mem-
bers of Ulohorus less resistant than Phil-
oponella. A large Zosis geniculatus (tra-
cheal pattern 2) population discovered in
Panama Canal Company warehouse B by
Yael Lubin thrives in the apparent ab-
sence of free water.

Tracheal patterns are usually consid-
ered to evolve from simple to more ex-
tensive patterns. However, the following
analysis of uloborid intrafamilial phylog-

Revision of Uloboridae • Opell 465

eny indicates that the more extensive tra-
cheal pattern 4 is either primitive in the
family or has evolved independently at
least five times. I favor the former expla-
nation because of its greater parsimony.
The literature, however, provides no in-
sight into selective pressures responsible
for reduction of the tracheal system and
seems to dismiss this possibility.

Apparent desiccation resistance of
PhiloponcUa and Zosis indicates that tra-
cheal patterns 1 and 2 are not detrimental
in this regard. Web-hanging postures and
relatively smaller femur diameters sug-
gest that members of these genera ex-
pend less energy during web monitoring
and prey entrapment and therefore may
require less oxygen. In this case a well-
developed tracheal system might prove
disadvantageous because it would re-
quire an added initial investment of tis-
sue and perhaps slightly raise basal met-
abolic rate and respiratory water loss.

Discussion. Members of the family
Uloboridae possess six potentially syn-
apomorphic (shared, uniquely derived)
characters: 1. loss of poison glands, 2.
presence of a ventral row of macrosetae
on the fourth metatarsae and tarsae (Plate
1-A, C), 3. two apical cymbial setae on
the male palpus, 4. production of an orb-
web or web derived from an orb, 5. pres-
ence of a row of trichobothria on the sec-
ond through fourth femora, and 6. stout
tracheal trunks which extend into the
cephalothorax. Absence of a plesio-
morphic character is generally not admis-
sible as phylogenetic evidence. How-
ever, the fact that all uloborid genera are
united by the five other characters listed
above and are the only araneomorph spi-
der genera to lack poison glands seems
good reason to consider this loss a char-
acter uniting all uloborids. This charac-
ter, however, offers little insight into
interfamilial relationship. Ventral macro-
setae occur on the metatarsi and tarsi
of other spiders, but those of uloborids
are larger, more perpendicular, and more
precisely in register than those present

in other spiders I have examined. These
macrosetae appear to handle silk used
to wrap prey (see Natural History) and,
as uloborids rely heavily on this abil-
ity, the extreme development of this
setal row may be considered a family
character. The structure of these macro-
setae distinguishes them from setae of
the theridiid comb row, also situated on
the fourth tarsus. Two prominent setae
on the apex of the male's palpal tarsus
(Plate 3-A, 6-C, D) appear homologous
with and derived from stridulatory picks
of Tangaroa, Waitkera, and Polenecia
palpi (Figs. 19, 28, 46). Until similar
stridulatory apparati or setae are discov-
ered in other spider groups, these setae
must be considered unique to the family
Uloboridae.

I interpret behavior responsible for
orb-web production as an apomorphic
(primitive) character of the family Ulo-
l)oridae and the behavior of Polenecia,
Hyptiotes, and Miagrammopes as a mod-
ification of this behavior. Several inge-
nious schemes have been proposed to
explain the monophyletic or (more often)
polyphylyetic origin of the orb-web from
various simple web types (e.g., Kaston,
1964, 1966; Kullmann, 1972), but these
remain largely untested by morphologi-
cal or behavioral evidence. This study is
not intended to be a comprehensive
treatment of interfamilial relationship,
rather it attempts to present a clearly de-
fined hypothesis of uloborid intrafamilial
relationship which will facilitate these
broader studies. Presence of femoral
trichobothria on the legs of all uloborids
(Fig. 206) as well as the legs of some ara-
neids provides support for the relation
of these groups and monophyletic evo-
lution of the orb-web. Uloboridae have a
single row of prolateral femoral tricho-
bothria on the third and fourth legs and
a single retrolateral row on the second
legs. The first femur has either a single
or double row of dorsal trichobothria. It
is generally recognized that members of
the genus Leucauge have two rows of

466 Bulletin Museum of Comparative Zuology, Vol. 148, No. 10

Table 2

Characters Used in Table 3

Character

Primitive State I I

Derived State a|

bE22

1 Calamisfrum length/ metatarsus IV

2 Embolus in cross section

3 Radix

4 Lengthi of male palpal femur

5 Conductor

6 Female genital proiection

7 Median apopfiysis

8 IVIale first femur

9 Anterior lateral eyes

10 Median spermatfieca

11 Macrosetae on cymbial margin

12 Accessory glands

13 Dorsal tnchobothria on Q femur I

14 Female genitalia

15 Embolus guide

16 Stridulatory apparatus

17 Posterior abdomenal extension

18 Apical cymbial spine

19 Anterior half of carapace

20 Posterior lateral eye tubercules

21 Large middle hemafodocha

22 Median apophysis

23 Male palpal tibia

24 Anterior median eyes

25Endite length

26 Cephalothoracic tracheal trunks

27 Setal tuft on female tibia I —

28 Tegular spur

29 Median apophysis spur

30 Female genital openings

31 Tracheoles

32 Epigynal atrium

Less than half

Flat

Absent

4 X diameter
Absent

Absent

Present
Straight

Present

Absent
Two

Absent

1 row

Haplogyne

Does not include radix

Present

Absent

Rounded

Absent

Does not form a bulb

Small

Present

One - and - one - third width
Two

Absent

Absent
Absent
Posterior

Pass into cephalothorax
Absent

a. half b. more than half

a. round b. crescent

a. present b. bifurcate

3 X diameter

Present

a. median b. lateral

Absent

With distal crook

Absent

Present

Three

Present

2 rows

Entelegyne

Includes radix

Absent

Present

Narrowed

Present

Forms a large bulb

With extension

Absent

Twice width

a. four b. absent

Present

a. large b. small

a. claw-like b. concave

Ventral

Restricted to abdomen

a. undivided b. divided

Revision of Uloboridae • OpcU

46'

Table 3. Cladogram of Uloboridae genera using characters described in Table 2. Boxes
marked with an x represent a character state reversal.

OCT = OCTONOBA
PURUM = PURUMITRA

468 Bulletin Museum uf Comparative Zoologij, Vol. 148, No. 10

prolateral trichobothria on the fourth pears derived from "a" and Character 26

femora. It this were the only araneid ge- where "a" and "b" appear independently

nus to have such trichobothria their pres- derived. The structure serving as an em-

ence might be explained by conver- bolus guide in haplogyne spiders appears

gence. However, the tetragnathid genus to be homologous with the median

A^riofiiuitha has a single row of tricho- apophysis oi Waitkem (Figs. 28, 29) and,

bothria on the dorsal surface of the first for this reason, I consider presence of a

femur, the dorsoretrolateral surface of the median apophysis a primitive character

second femur, and the prolateral surfaces of uloborids and its absence in Tangaroa

of the third through fourth femora. The a derived condition.

Metinae genus AlcimospJienus and Leu- Tangaroa is distinguished by derived

cauge argijra (the latter previously placed states of Characters 7, 8, and 9 and the

in the Metinae genus Plesiometa) have remaining fifteen genera by derived

dorsal trichobothria on the first femur and states of Characters 1 through 6. Waitkera

two rows of trichobothria on the dorsal is the only genus with derived states of

surface of the second femur and the pro- Characters 10 and 11 and is united with

lateral surfaces of the third through the Polenecia, Hijptiotes, Siratoba, and Ar-

fourth femurs. /sfoH by the derived state of Character 12.

Based on the synapomorphic charac- The remaining ten genera share derived

ters of orb-web production and the pres- states of Characters 13 and 14. Miagram-

ence of rows of femoral trichobothria, the mopes, Sijbota, and Orinomana are

family Araneidae appears to be the only united by the derived state of Character

family with which Uloboridae may be 20 and the remaining seven genera by

objectively allied. This hypothesis may derived states of Characters 6, 21, 22, and

be shown false by discovery that these 29. Uloborus is characterized by the de-

trichobothrial rows are not homologous rived state of Character 27 and the re-

or that they are shared by members of maining six genera by Character 26-b.

other families. Zosis, Furumitra and Octonoba share

The extensive tracheal pattern which the derived state of Character 28 and the
I interpret as primitive within the Ulo- remaining three genera share the derived
boridae is also found in members of other state of Character 30. Other shared, de-
families and, for this reason, cannot at rived characters further define the clado-
this time be regarded as a synapomorphic gram presented in Table 3.
character of uloborids. However, this Conclusions on relationships of ulob-
character does separate Uloboridae from orid genera (Table 3) differ from tradi-
Dinopidae which have a four-branched tional division of the family into three
tracheal system restricted to the abdo- subfamilies and more precisely place the
men. genera. I have chosen not to present a

Most characters (Table 2) used in the formal classification for the family Ulo-

chidistic analysis (Table 3) of generic re- l)oridae at this time. I do so not out of

lationship are discussed above and can disagreement with arguments favoring a

also i)e found in genus descriptions. In well-defined and detailed classification,

the seven cases where two derived states but out of the belief that this investiga-

are designated for a character each state tion will stimulate additional studies

is treated as an alternate possibility and which will test and improve this hypoth-

designation of a state as "b" in Table 2 esis of intrafamilial relationship. Because

does not imply that it was derived from a cladogram is translated directly into a

state "a." Ensuing analysis of this data cladistic classification, lack of a hierar-

can provide some directional evidence as chical scheme does not prevent a clear

in the case of Character 6 where "b" ap- understanding of the underlying hypoth-

Revision of Ulobofudae • Opell 469

esis of relationship nor preclude its test-
ing.

Uloboridae Thorell

Uloborinae Thorell, 1869, N. Act. Reg. Soc. Sci.,
Upsalaiensi.s, (.3)7: 1-108 (In Epeiroidae). Type
genus Ulohorus Latreille, 1806.

Uloboridae: — O. Pickard-Camhridge, 1870, Zool.
Rec, 7: 207-224.

Diagnosis . Members of the family
Uloboridae are distinguished from all
other spiders by the lack of poison
glands. They are the only cribellate spi-
ders which spin an orb-web, have a row
of trichobothria on femora II through IV
(Fig. 206), and have a prominent ventral
row of macrosetae beginning on the dis-
tal half to third of metatarsus IV and ex-
tending nearlv the full length of tarsus IV
(Plate 1-A, C)'.

Description. The carapace of most
uloborids is oval (Figs. 13, 23, 72) to pear-
shaped (Figs. 40, 132, 162), but that of
Hyptiotes is anteriorly constricted (Fig.
55) and that of Miagrammopes nearly
rectangular (Figs. 78, 86). In males of
some genera the anterior median eyes are
situated on a small median mound (Figs.
14, 193). The thoracic groove of females
and in many genera of males is a shallow
pit (Figs. 13, 14, 40). Males of Zosis, Po-
nella, and PhiloponeUa have a broad,
transverse thoracic groove (Figs. 163,
193, 217). All eyes are nearly equal in
size and lack a tapetum. In members of
the genus Tangaroa anterior lateral eyes
are reduced to small pigment spots (Figs.
13, 14) and in members of the genus Mia-
grammopes the anterior eyes are absent
(Figs. 78, 80). The eight eyes of the re-
maining 14 genera are arranged in two
rows. When viewed from above the an-
terior row is procurved in Polenecia (Fig.
50), straight in Hyptiotes (Fig. 55) and
recurved in the remaining genera (Figs.
13, 23, 120). The posterior eye row is
nearly straight in PhiloponeUa (Figs. 204,
205) and in some Miagrammopes (Figs.
78, 80) and recurved in the remaining
genera (Figs. 13, 23, 55). Posterior lateral

eyes of Hyptiotes, Syhota, Orinomana,
and Miagrammopes are on conspicuous
lateral tubercles (Figs. 55, 78, 79, 100,
121). The sternum of most Miagram-
mopes species is divided (Figs. 81, 82,
92, 93) and in the remaining genera en-
tire, being widest between the first and
second coxae (Fig. 164). The labium is
not rebordered and, although free from
the sternum (Figs. 81, 82, 164), its sepa-
ration from the latter is not easily dis-
cernible in most genera. Endites of Mia-
grammopes species are about twice as
long as wide and in others about one-and-
one-third times as long as wide. Endite
serrulae of all genera are formed by a sin-
gle row of closely spaced denticles (Plate
7-A). Chelicerae lack a prominent boss
and their prolateral and retrolateral fang
furrow margins each have a cluster of
many small teeth and in some genera one
or more larger teeth. Each fong's median
surface has a row of small denticles
which runs nearly its entire length. Its tip
lacks a poison opening. Palpal tarsus of
immatures and females each bears an api-
cal serrate claw. The first leg tarsal organ
of female representatives of all genera
except Tangaroa, Siratoba, Orinomana,
Purumitra, and Daramuliana was exam-
ined with the scanning electron micro-
scope and found to be dome-shaped with
a central pit (Plate 2-D). The first femur
has dorsal trichobothria arranged in one
or two rows (Table 3). The second femur
has a single row of trichobothria which is
proximally dorsal, becoming distally ret-
rolateral (Fig. 206). The third and fourth
each have a single row of trichobothria
which is proximally dorsal and distally
prolateral. Plumose setae are present on
legs. Each leg has three tarsal claws, the
dorsal two being serrate. Leg formula
(longest to shortest): I, IV, II, III. Paired
patellar lyriform organs (Plate 2-C) are
retrolateral and situated at the proximal
margin of a broad, open, weakly sclero-
tized notch (Figs. 58, 84, 166). In Plate
2-C this weakly sclerotized notch is rec-
ognized by the absence of setal sockets.

470 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

The fourth metatarsus is Literally com-
pressed in the proximal region which
bears the dorsal calamistrum (Plate 1-B,
C). Although only one row of calamistral
setae is well developed, a prolateral row
of short setae (Plate 1-B) adjacent to the
primary row betrays the double nature of
the calamistrum. A row of stout, nearly
perpendicular, ventral setae each with a
spiraled shaft and small, hooked tip
(Plate 1-A) extends from the center or dis-
tal third of the fourth metatarsus to the
tip of tarsus IV. Presumal)ly, these setae
serve to handle silk during prey wrap-
ping (see Natiual History). The pedicel's
lorum (Fig. 168) consists of a broad, an-
terior, shield-shaped plate, a smaller,
posterior, H-shaped plate, and a pair of
small lateral plates each bearing a lyri-
form organ (not shown). Abdomen shape
is quite variable and ranges from oval
(Figs. 24, 40, 42, 56) to peaked (Figs. 74,
122) to cylindrical (Figs. 78, 86). Abdo-
men of Polenecia and Sybota have a nar-
row posterior extension (Figs. 51, 114,
116). The four dorsal humps present in
some genera (Figs. 56, 148, 149) appear
to correspond to attachment points of the
four pairs of dorsal-ventral abdominal
muscles. In many genera only one pair of
humps is visible (Figs. 120, 122, 132, 195,
260). The heart has three pairs of ostia
which in light colored specimens can be
seen through the integument (Figs. 40,
98, 133, 159, 195). The cribellum is un-
divided and bordered anteriorly by a
weakly sclerotized region which bears
fine setae (Plate 1-D). Cribellum and cal-
amistrum are absent in second in star spi-
derlings when they emerge from the egg-
sac as well as in mature males. In males
the area previously occupied by the cri-
bellum does not bear a colulus, but has
a weakly sclerotized integument similar
to that present anterior to a functioning
cribellum. Just anterior to the cribellum
is a single median spiracle whose broad,
shallow atrium gives rise to a pair of lat-
eral chambers, each giving rise to a stout
anterior tnnik and several smaller pos-

terolateral tracheae (Figs. 1, 2, 3, 5, 7, 9,
11). A more extensive discussion of the
tracheal system is found under Intrafa-
milial Phylogeny. Anterior spinnerets ap-
pear to have three segments: a large basal
segment; a very short, ringlike middle
segment; and a short, dome-shaped api-
cal segment. Median spinnerets have a
single segment and posterior spinnerets
have two cylindrical segments, the prox-
imal one being longer. The anal tubercle
has two segments. A detailed discussion
of male and female genitalia appears un-
der Intrafamilial Phylogeny and Map 1
summarizes the distribution of the fami-
ly's 16 known genera.

Note. Type specimens of Uloborus
minutus Mello-Leitao and U. tetramac-
ulatus Mello-Leitao were not examined
during the course of this study. Types of
v. coUinus and V . trillineatus described
by Keyserling should be housed in Pol-
ska Akademia Nauk Instytut Zoologiez-
ny, Warsaw, Poland, but appear to have
been lost. Uloborus formosus Marx (in
Banks) was destroyed in the 1906 Cali-
fornia earthquake. The type specimen of
U. niger Mello-Leitao belongs to the fam-
ily Dictynidae and the type specimen of
U. ater Mello-Leitao is an immature
male. Uloborus sexmucronatus Simon,
the type and only known species of
Lehtinen's genus Astavakra should be
housed in the Museum National d'His-
toire Naturelle, Paris, but could not be
located. Conseciuently, this is the only
described genus of Uloboridae not treat-
ed in this study.

Generic Key for Uloborid Males
(Males oi Orinomana are unknown)

1. Posterior lateral eyes (PLE) on prominent

lateral tubercles (Figs. 55, 80, 103) 2

- PLE not on lateral tubercles 4

2(1). Anterior eyes absent (Figs. 80, 87); en-
dites and labium at least twice as long as

wide (Figs. 82, 92); Cosmotropical

Miagrammopes

Anterior eyes present (Figs. 55, 103); en-
dites and labium length equal to or less
than 1.3 width 3

3(2). Anterior half of carapace conspicuously

Revision of Uloboridae • Opell 471

DARAMIILIANA SAMOA and VITI

LEVU ISLANDS

• —LOCALITY RECORD
R— PROBABLE GENUS RANGE

Map 1. Distribution of uloborid genera.

narrowed (Fig. 55); in dorsal view ante-
rior median eyes (AME) separated from
anterior carapace edge l^y at least two eye
diameters (Fig. 55); length of femur I
equal to carapace length; median ocular -

quadrangle four times as wide posteriorly
as anteriorly; Holarctic and Oriental
(Map 1) Hyptiotes

- Carapace oval (Fig. 103); in dorsal view

AME at anterior edge of carapace (Fig. 7(6).

103); length of first femur 1.5 to 1.7 times
carapace length; median ocular quadran-
gle 1.5 to 2.0 times as wide posteriorly as
anteriorly; western South America (Map

1) Sybota

4(1). Anterior median eyes (AME) on a con-
spicuous anterior projection (Fig. 14); —
ALE absent, represented only by small,
dark spots; first femur with a distal crook
(Fig. 21); South Pacific islands (Map 1)
Tangaroa

- AME not on a conspicuous projection;
ALE present; femur I without a distal

crook 5 8(7).

5(4). Palpus with a large, dome-shaped median
apophysis bulb and a small, claw-shaped
median apophysis spur (Plate 7-A-C) __ 6

- Palpus otherwise; median apophysis
bulb flattened (Plates 6-C, D, 7-D; Figs.
28, 46, 47, 157), reduced (Plate 3-C, D;

Figs. 68, 69), or median apophysis is a -

long projection (Plate 3-A, B) 9

6(5). Palpus without a conductor and with a

broad, flat tegular spur serving as an em-
bolus guide (Plate 7-C); first femur (Fig.
167) witli one prolateral and three dorsal

macrosetae; cosmotropical Zosis

Palpus with a conductor and without a
tegular spur (Plate 7-A, B; Figs. 201, 218,
253, 270); first femur with two or three
prolateral, one or two retrolateral, and

two dorsal macrosetae (Fig. 202) 7

Palpus with a large conductor extending
along the distal embolic surface and hav-
ing a long, thin projection at its base (Fig.
201); first tibia wi'h three prolateral,
three retrolateral, and four dorsal macro-
setae (Fig. 202); southeastern South

America (Map 1) Ponella

Palpus either without a conductor along
the distal embolic surface (Plate 7-A) or
if conductor is longer it consists of a basal
lobe and a long conductor spur (Plate
7-B); first tibia with at least four prolat-
eral, four retrolateral, and six distal mac-
rosetae 8

Carapace oval with a broad, transverse tho-
racic groove (Fig. 217); posterior eye row
nearly straight; clypeus height in anterior
view 0.7 to 1.5 AME diameter; conductor
spur present (Plates 7-B; Figs. 218, 236);
first femur without numerous trichoboth-

ria; cosmotropical Philoponella

Carapace pear-shaped with conspicuous-
ly narrowed cephalic region and a narrow
thoracic depression (Figs. 133, 142); pos-

472 Bulletin Museum of Comparative Zoolo^ij, Vol. 148, No. 10

teiior e\e row recim'ed such that a hue
touching PMEs' posterior margins passes
anterior to the PLEs l)y half a PLE di-
ameter; clypeus height half an AME di-
ameter; conductor spur absent (Plate 7-A;
Figs. 135, 143); first femur with numer-
ous trich()i)()thria (Fig. 131); cosmo-poli-

tan Uloborus

9(5). Median apophysis spur a broad, concave
or enrolled sclerite (Plates 6-C, D; 7-D);
tegular spur usually present (Plates 6-C,
7-D) _'_ 10

- Palpus otherwise 11

10(9). Male carapace length greater than 1.4

mm; length of first femur 1.5 to 1.7 that
of carapace; median apophysis bulb re-
duced, median apophysis spin" concave
(Plates 6-C, D, 7-D); southeastern United
States, China and adjacent Pacific islands
(Map 1) Octonoha

- Carapace length less than 0.9 mm; length
of first femur twice that of carapace; me-
dian apophysis bulb large and laterally
flattened; median apophysis spur broad
with a central depression (Fig. 157); Pa-
cific islands (Map 1) Purumitra

11(9). Embolus short, pointed, and free from
guide for most of its length (Figs. 28, 29,
46, 47); c>nil)iiun with two or three short,
stout apical macrosetae; each endite's lat-
eral surface with a lobe bearing stridula-

tory ridges (Figs. 28, 46, 47) 12

Embolus either broad and flat (Plate 3-A
B) or resting in a grooved radi.x (Plate
3-C, D; Figs. 68, 69); cymbium without
such spines; endite lacks a lateral lobe
and stridulatory file 13

12(11). Anterior eye row recui-\'ed (Fig. 23); first
tibia with six prolateral, two retrolateral,
six dorsal, and no ventral macrosetae
(Fig. 27); cymbium with three sto>it distal
marginal macrosetae (Fig. 28); palpus
with a pointed, grooved median apophy-
sis serving as an embolus guide and with
a long thin radix (Figs. 28, 29); New Zea-
land (Map 1) Waitkeru

- Anterior eye row procurved (Fig. .50);
first tibia with two prolateral, two retro-
lateral, one dorsal, and six ventral mac-
rosetae (Fig. 49); cymbium with two stout
distal, marginal macrosetae and a thin
distal cymbial extension (Figs. 46, 47);
palpus with a thin proximal median
apophysis lobe and a broad flat conductor
with a thin apical lobe together as a
guide; radix not conspicuous (Fig. 46,
47); Mediterranean (Map 1) Polenecia

13(11). Radix a thin, broad sclerite, bifurcate at
its tip and not surrounding embolus
(Plate 3-A, B); PME at least twice as far
from one another as from PLE (Fig. 41);
total length about 1.4 mm; southern Mex-

ico through Central America (Map 1)

Ariston

Radix a large, coiled sclerite whose
grooved inner surface holds the embolus
(Plate 3-C, D; Figs. 68, 69); PME 1.0 to
1.5 times as far from one another as from
the PLE (Fig. 72); total length about 2.7
mm; southwestern United States and
central Mexico (Map 1) Siratoba

Generic Key for Uloborid Females

1. Abdomen with a prominent posterior pro-
jection extending beyond spinnerets
(Figs. 51, 102, 116) 2

- Abdomen without such a posterior pro-
jection (Figs. 15, 24, 56, 74, 122, 197) __ 3

2(1). Posterior lateral eyes (PLE) near cara-
pace border on prominent lateral tuber-
cles (Figs. 100, 115); anterior eye row re-
curved when viewed dorsally; epigynum
without a midventral posterior projec-
tion (Figs. 105, 112, 117); Chile (Map 1)
Sybota

- PLE not near carapace border and not on
lateral tubercles (Fig. 50); anterior eye
row procurved when viewed dorsally;
epigynum with a midventral posterior
projection (Figs. 52, 53); Mediterranean
(Map 1) Polenecia

3(1). Anterior eyes lacking (Fig. 78); endites
and labium at least 1.5 times as long as

wide (Figs. 81, 92); cosmotropical

Miiifirammopes

- Anterior eyes present, although anterior
lateral eyes (ALE) may be reduced;
length of endites and labium 1.3 or less
their width (Fig. 164) 4

4(3). ALE reduced, represented only by dark
pigment spots (Fig. 13); genital area
when viewed ventrally a low, weakly
sclerotized mound (Fig. 16); South Pacif-
ic islands (Map 1) Tan'^aroa

- ALE present; genital area either weakly
sclerotized with a central peak and a pos-
teromedian groove (Fig. .30) or distinctly
modified and (usuallv) well sclerotized
(Figs. 32, 59, 65, 169, 178, 198, 208, 236) 5

5(4). Posterior lateral eyes (PLE) on distinct

lateral tubercles (Figs. 23, 55, 121) 6

- PLE not on distinct lateral tubercles
(Figs. 72, 175, 186, 192) 8

6(5). Anterior half of carapace conspicuously
narrower that posterior half (Fig. 55); an-
terior eye row straight; median ocular
area's anterior border one-fourth the
width of its posterior border; length of
first femur less than that of carapace; Hol-
arctic; Oriental (Map 1) Hyptiotes

- Carapace oval or pear-shaped (Figs. 23,
121); anterior eye row recurved; median

Revision of Uloboridae • Opell 473

ocular area's anterior border three-fifths
to fi\'e-si\ths as wide as its posterior bor-
der; length of first femur at least 1.4 that
of carapace 7

7(6). Anterior eye row recurved such that a
line across the anterior median eyes'
(AMEs') posterior margins passes along
the .ALEs' anterior border (Fig. 23); in
dorsal view the clypeus does not project
anteriorh'; abdomen rounded in lateral
view (Fig. 24); genital area a weakly
sclerotized mound with a central peak
and a posteromedian notch (Fig. .30);
New Zealand (Map 1) Waitkcra

- Anterior eye row recurved such that a
line across the AMEs' posterior margins
passes through the anterior third of .\LE
(Fig. 121); in dorsal view the clypeus ex-
tends anteriorly a distance equal to one
AME diameter; abdomen peaked in lat-
eral view (Fig. 122); epigynum well
sclerotized, flattened, and with a poste-
rior indentation (Figs. 123, 126); western
South America (Map 1) Orinomana

8(4). Epigvnum with a pair of posterolateral

lobes (Figs. 1,37, 154, 178, 184, 198) __._ 9

- Epigynum with one or more ventral atria
(Figs. 160, 189, 208, 239, 250) or with a
posteriorlv directed median projection
(Figs. .32, '.36, 43, 65, 75) J 12

9(8). Posterior eye row recurved such that a
line across the posterior median eyes'
(PMEs') posterior margins passes along
or anterior to the PLEs' anterior margins
(Figs. 132, 175); posterior epigynal lobes
slender and their bases contiguous (Figs.
137, 1.54, 178, 184) . 10

- Posterior eye recurved so that such a line
passes through the center of the PLEs
(Figs. 162, 192); posterior epigynal lobes
broad and their bases separated by a dis-
tance equal to the width of each (Figs.
169-171, 198, 199) _. 11

10(9). Posterior eye row recurved such that a
line across the PMEs' posterior margins
passes along the PLEs' anterior margins
(Fig. 175); width of cephalic region at the
ALEs two-thirds the maximum carapace
width (Fig. 175); median ocular area's an-
terior width and length 0.7 its posterior
width; epigynum's posterior surface with
a conspicuous atrium or pair of lateral
atria (Figs. 179, 185); United States and
China and adjacent Pacific islands (Map
1) Octonoba

- Posterior eye row recurved so that such
a line passes anterior to the PLEs by a
distance equal to at least one-fourth a
PLE diameter (Figs. 132, 148); cephalic
region at ALE half the carapace width;
median ocular area square; epigynum's
posterior surface without conspicuous

atria (Figs. 1.38, 146); cosmopolitan

Uloborus

11(9). Epigynum with two low, rounded pos-
terolateral mounds anterior to which the
openings are found (Figs. 198, 199); total
length less than 4 mm, southeastern
South America (Map 1) Ponelhi

- Epigynum with two flattened, ventrally
or posteriorly directed lobes whose ven-
trolateral surfaces each bear a triangular
sclerite and whose posterior surfaces
form lateral atria in which openings are
found (Figs. 169-171); total length great-
er than 6 mm; cosmotropical Zosis

12(8). Epigynum with a single median, ventral
atriinn (Figs. 32, 36, 43, 65, 75, 208, 239,
2.50, 266, 27,5) 13

- Epigynum with two or more atrial divi-
sions (Figs. 160, 189) 15

1,3(12). Epigynum with a ventral atrium (Figs.
208, 239, 2,50, 266, 275); total length 2.9
mm or greater; posterior eyes aI)out
equally spaced (Figs. 216, 260); cosmo-
tropical Philoponella

- Epigynum with a ventromedian projec-
tion extending over a posterior atrium
(Figs. 32, 43, 65, 75) or if median projec-
tion is less distinct (Fig. 36) total length
2.5 mm or less and PME three times as
far from one another as from PLE (Figs.
40, 41) 14

14(13). Carapace length and width equal (Fig.
41); PME 2.5 to 3.0 times further from
one another than from PLE (Figs. 40, 41);
clypeus height in anterior view half or
less an AME diameter; total length less
than 2. ,5 mm; epigynum without a distinct
posterior plate (Figs. 33, 37, 44); southern
Mexico through Central America (Map 1)
Ariston

- Carapace 0.8 times as wide as long (Fig.
72); posterior eyes about equally spaced
(Figs. 71, 72); clypeus height 0.8 to 1.0
AME diameter, total length 3.0 mm or
greater; epigynum with a distinct poste-
rior plate (Figs. 66, 76); southwestern
United States and central Mexico (Map
1) Siratoba

15(12). Epigynum with two lateral atria (Fig.
189); median ocular area's anterior width
half its posterior width (Fig. 186); cara-
pace three-fourths as wide as long; total
length 4.0 to 4.6 mm; Australia and Pa-
cific islands (Map 1) DaramuUana

- Epigynum with an anterior and posterior
median atrium, each divided by a median
septum, and a pair of anterior and poste-
rior lateral atria (Fig. 160); median ocular
area nearly square (Fig. 1,59); carapace
width and length equal; total length

about 3.0 mm; Pacific islands (Map 1)

Purumitra

474

Bulletin Museum of Cumparative Zoology, Vol. 148, No. 10

Tangaroa Lehtinen
Figures 13-22, Map 1

Tangaroa Lehtinen, 1967, Ann. Zool. Fennici, 4:
199-468. Type species by original designation
Ulohonis tahiticnsis Berland, 1934, Ann. Soc.
ent. France, 103: 321-336. Male holotype and 3
female paratypes in Museum National d'Histoire
Natiirelle, Paris, examined. The genus name is
feminine.

Diagnosis. Males and females are dis-
tinguished from those of all other genera
by lack of ALE (Figs. 13, 14). Males also
differ from those of other genera by pres-
ence of: 1. AME on an anterior projection
(Fig. 14), 2. a distal crook in femur I (Fig.
21), and 3. a palpus with a folded cymbial
tip and only a flattened embolus (Fig. 19).
Females are also differentiated by weak-
ly sclerotized genitalia in the form of a
low mound (Figs. 16, 17).

Description of Type Species. Total
length of female 4.6 to 5.6 mm. Carapace
0.85 times as wide as long. In lateral view
cephalic region rises slightly from shal-
low thoracic depression to PME. In fe-
males it then slopes downward and in
males upward to AME. Thoracic region
slopes gradually to the petiole. Male and
female ALE absent, represented only by
darkly pigmented areas (Figs. 13, 14). At
AME carapace width 0.35 its maximum
width. Male AME on conspicuous,
anterodorsal projection (Fig. 14). Poste-
rior eye row recurved such that line
across posterior margins of PME passes
anterior to PLE. Median ocular area in
females nearly square (slightly wider
than long) (Fig. 13) and in males trape-
zoidal, anterior edge slightly wider than
posterior edge and length slightly less
than posterior width (Fig. 14). In anterior
view clypeus height of females equal to
AME diameter and of males 2.0 to 2.5

times AME diameter. Female sternum
two-thirds and male sternum four-fifths
as wide as long. Female palpal femur lat-
erally flattened and calamistrum one-
third as long as metatarsus IV. Female
first femur 2.1 times and male first femur
1.9 times carapace length. Male first fe-
mur bears four dorsal, four prolateral, and
three retrolateral macrosetae. Tibia I
bears seven dorsal, eight ventral, five
prolateral, and five retrolateral macrose-
tae. Distal third of the male first tibia
forms a crook whose concave ventral sur-
face bears stout macrosetae at its poste-
rior margin (Fig. 21). Abdomen width
and height about half its length (^Fig. 15).
Two stout tracheal trunks extend into
cephalothorax.

Male palpus. Endite with a small, lat-
eral stridulatory file just dorsal to serrate
edge (Fig. 19). Two macrosetae on distal,
median surface of cymbium serve as
picks. Femur lacks ventral tubercles.
Cymbium large, its broad distal portion
folded laterally, apparently serving as
guide for embolus. Within this broad fold
and near the embolus' base is a dark, oval
region which close examination shows to
be part of the heavily sclerotized sperm
duct and not a median apophysis (Fig. 19,
20). No middle hematodocha is present.
Unlike other uloborids, the embolus is
flattened and is not closely associated
with a guide.

Genital Region. In ventral view this re-
gion appears as a lightly sclerotized, con-
vex shield (Fig. 16). In posterior view its
central, unsclerotized region has two me-
dian openings (Fig. 17), each leading to
a pair of lobed spermathecae (Fig. 18).
No fertilization duct leads from either
spermatheca.

Natural History and Distribution. In

Figures 13-22. Tangaroa tahitiensis (Berland). 13. Dorsal view of female carapace. 14. Dorsal view of male carapace.
15. Lateral view of female abdomen. 16. Ventral view of female genitalia. 17. Posterior view of female genitalia. 18. Dorsal
view of cleared female genitalia. 19. Retrolateral view of left male palpus. 20. Retrolateral view of embolus (black), sperm
reservoir, and fundus. 21. Prolateral view of male left, first femur, patella, and tibia. 22. Dorsal view of female rigfit first
femur showmg tnchobothria. Figures 23-27. Waitkera waltkerensis (Chamberlain). 23. Dorsal view of female carapace.
24. Lateral view of female abdomen. 25. Lateral view of female carapace. 26. Dorsal view of female right first femur. 27.
Prolateral view of male left first femur, patella, and tibia.

Revision of Uloboridae • Opell 475

-..^!;,'.-,vjK-.r'i':i.S)^.^-.-:'*-.L--'fi-i:,h-)'..t!-

^;;^

:)»...

RjS;:SA^^j^.^<5^s;-»va^;^&g-;.'^; ,

'^-^f^^aSf^-vy^^-^'a

476

Bulletin Musvudi of Comparative Zoology, Vol. 148, No. 10

addition to the type species found on Ta-
hiti in the Society Islands and Rapa in
the Tu])uai Islands, this uenus contains
Tan^arod (Ulohorus) dissiniild (Berland,
1924), found on New Caledonia and the
New Hebrides island of Epi (Map 1).
Wel)s of neither species are known.

Waitkera new genus
Figures 23-31, Map 1

Tan^arod [part]: — Lehtinen, 1967, Ann. Zool. Fen-
nici, 4: 199-468.

Type. Type species Vlohorus nditkcrcnsis Cliani-
'berlain, 1946, Ree. Auckland Inst. Mus., 3(2): 85-
97. Female holotype and three paratypes from
Watershed Reservoir, W'aitker Hills, Auckland,
New Zealand collected by C Chamberlain. Ho-
lotype and one paratype in G. Chamberlain col-
lection, two paratypes in Auckland Museum. The
genus name is feminine and is derived from the
locality of the type species.

Diagtiusi.s. Like Sybota and Orino-
mana, Waitkera females have PLE on
distinct lateral tubercles (Fig. 23). Unlike
Sybota (Fi.iis. 102, 114, 116) the abdomen
of Waitkera (Fig. 24) does not have a
prominent posterior extension and imlike
Orinomana (Figs. 120, 122), it is not
peaked. Posterior eye row of Waitkera
males and females only slightly recurved
such that a line across PMEs' posterior
margins passes through PLEs' centers
(Fig. 23). In Sybota (Figs. 100, 115) and
Orinomana (Fig. 121) (males of the latter
are unknown) such a line passes along
PLEs' anterior borders or through ante-
rior one-fourth of their diameters. First
femur of Waitkera males and females
twice as long as carapace, in Sybota and
Orinomana 1.3 to 1.7 times carapace
length. Genitalic region of Waitkera fe-
males (Fig. 30) a weakly sclerotized
mound, that of Sybota (Figs. 105, 112,
117) and Orinomana (Figs. 123, 126)
well sclerotized. Male palpus of Wait-

kera (Figs. 28, 29) has flattened, pointed
median apophysis serving as embolus
guide; prominent radix, and three short,
stout marginal cymbial macrosetae. Pal-
pus of Sybota (Plate 6-A, B) has well-de-
veloped conductor, two- or three-pronged
median apophysis, and lacks radix.

Description. Male total length 2.9
mm, carapace length 1.1 mm, and ster-
num length 0.7 mm. Female total length
3.5 mm, carapace length 1.1 mm, and
sternum length 0.7 mm. Carapace width
equal to length. Cephalic region rises
slightly from thoracic depression and tho-
racic region slopes gradually to petiole
(Fig. 25). At ALE cephalic region three-
fifths carapace width (Fig. 23). Both eye
rows recurved, posterior such that a line
across posterior margins of PME passes
through center of PLE. Lateral eyes on
slightly raised tubercles. Median ocular
area five-sixths as wide anteriorly as pos-
teriorly and as long as its anterior width.
Clypeus height in anterior view equal to
one AME diameter. Sternum 0.64 as
wide as long in females and 0.71 in
males. Female palpal femur not laterally
compressed. Female first femur 1.9 times
carapace length, male first femur 2.1
times carapace length. Female calamis-
trum half as long as metatarsus IV. Male
first femur with two dorsal, four prolat-
eral, and three retrolateral macrosetae
(Fig. 27). First tibia with six dorsal, six
prolate ral, and two retrolateral macrose-
tae. Female abdomen two-thirds as wide
and five-sixths as high as long (Fig. 24);
maximimi width and height attained half
way along length. Male abdomen cylin-
drical, half as wide and high as long. Car-
apace gray with broad light lateral mar-
gins and narrow median stripe extending
from thoracic pit through median ocular
area. Endites, labium, and sternum gray,

Figures 28-31. Waitkera waitkerensis (Chamberlain). 28. Retrolateral view of left male palpus. 29. Dorsolateral view of
distal region of left male palpus. 30. Ventral view of female genitalia. 31. Dorsal view of cleared female genitalia. Figures
32-34. Ariston albicans (O. Pickard-Cambridge). 32. Ventral view of female genitalia. 33. Posterior view of female genitalia.
34. Dorsal view of cleared female genitalia. 35. Dorsal view of first female femur. Figures 36-38. Ariston mazolus n. sp.
36. Ventral view of female genitalia. 37. Posterior view of female genitalia. 38. Dorsal view of cleared female genitalia.

Revision of Uloboridae • Opell All

~^-^Ai Ja

478 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

the latter with narrow white median ard-Cambridge, 1896, ;/;/(/. The generic name is

rr-.i ■ -.i • 1 1 •. masculine.
stripe. Iiliiae gray with wide white prox-
imal and median rings. Dorsum of abdo- Note. The small, generalized ulobo-
men white with five to six posteromedian rids which have been placed in Ariston
gray chevrons. Sides with transverse, appear to form two distinct genera, sep-
light gray stripes. Venter of abdomen arated l)y differences in eye arrangement
gray with white book lung covers, white and features of male and female genitalia,
paraxial stripes just behind epigastric fur- In addition to A. albicans the now-re-
row, and three pairs of white spots just stricted genus Ariston contains two new
anterior to cribellum. Two stout tracheal species, A. aristus and A. mazolus. Ar-
trunks extend into cephalothorax. iston referens and a new species have

Male Palpus. Lateral lobe on each en- been placed into the new genus Sira-

dite bears stridvilatory ridges (Fig. 28). toba.

Three stout macrosetae on posterodistal Diagnosis. Ariston is the only known

cymbial rim serve as picks. There are no uloborid genus with mature females hav-

ventral femoral tubercles or middle he- ing a total length of less than 2.5 mm and,

matodocha. Short, curved embolus lies in besides Siratoba (Figs. 65, 75), the only

trough on posterolateral face of flat, distal American genus whose members have a

median apophysis lobe (Figs. 28, 29). posterior epigynal hood (Figs. 32, 36, 43).

Smaller, proximal median apophysis lobe In Ariston the posterior atrium formed

lies between distal lobe and embolus by this hood lacks the prominent poste-

base. rior plate found in members of Siratoba

Female Genitalia. Genitalic area not (Figs. 66, 76). Ariston is found from

heavily sclerotized (Fig. 30), consisting southern Mexico through Central Amer-

of ventral mound with central peak and j^a and Siratoba has been collected only

posterior notch. Chamberlain (1946) de- f^om southwestern United States and

scribes an epigynal opening at each dor- central Mexico. In Ariston (as compared

sal margin of this notch, but these appear ^jth Siratoba) females, PMEs separated

to be openings of small, blind lateral by a distance at least two (Fig. 41) (in

bulbs (Fig. 31). Large, blind, spherical, Siratoba no more than one-and-one-half,

median spermatheca opens only into the Yig. 72) times that separating AMEs,

vagina through broad, hood-shaped pme separation two-and-one-half to

opening. Implications of this haplogyne three (one-and-one-half in Siratoba)

(sensus VViehle, 1967) condition are dis- times that of the PME-PLE separation,

cussed under phylogeny. PME and PLE mounds are confluent (in

Natural History and Distribution. The Siratoba separate) in anterior view, clyp-

single known species in this genus con- eus height equal to or less than half (in

structs small orb-webs among shrubs and SiratoI)a four-fifths to one) AME diame-

grass (Forster, 1967) and is known only ter, carapace length and width about

from New Zealand (Map 1). equal (in Siratoba carapace 0.80 as wide

as long), first femur 1.8-2.0 (in Siratoba

Ariston O. Pickard-Cambridge 1.5) times carapace length, and abdomen

Figures 32-45, Plate 3-A, B; Map 1 oval (Figs. 40, 42) (in Siratoba with mid-

Aristun O. Pickard-Cambndge, 1896, Bio. Cent.- ^le peak, FigS. 71, 74), three-fourths (in

Amer., Zool., Arachnida, 1: 161-224. Type Siratoba half) as Wide and three-fourths

species by monotype Ariston dlhicans O. Pick- as high as long. The only kllOWn males

Figure 39. Ariston mazolus n. sp., prolateral view of male left first femur, patella, and tibia. Figures 40-45. Ariston
aristus n. sp. 40. Dorsal view of female. 41. Female carapace. 42. Lateral view of female abdomen. 43. Ventral view of
female genitalia. 44. Posterior view of female genitalia. 45. Dorsal view of cleared female genitalia. Figures 46-49.

Revision of Uloboridae • Opell 479

Polenecia products (Simon). 46. Retrolateral view of left male palpus. 47. Prolateral view of left male palpus. 48. Retrolateral
view of embolus (black), sperm reservoir, and fundus. 49. Prolateral view of male, left, first femur, patella and tibia.

480 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

belong to A. mazolus and are distin-
guished as the smallest adult male ulo-
borids known (total length less than 1.4
mm) and the only known uloborid males
to have a series of six ventroprolateral
maerosetae on femur I (Fig. 39), a bifur-
cate radix (Plate 3-A, B), and a short, flat
eml)olus with a blunt tip.

Description. Carapace width and
length ecjual. Cephalic region rises
slightl)' from shallow thoracic depres-
sion, at level of ALE is about 0.6 carapace
width. Both eye rows about equally re-
curved such that a line across posterior
margin of median eyes passes along an-
terior margin of lateral eyes (Fig. 41).
Median ocular area about as wide ante-
riorly as posteriorly and about four-fifths
as long as its anterior width. In anterior
view, clypeus height of females half or
less AME diameter and in males one-
and-one-half times AME diameter. Ster-
num width about 0.68 length. Female
palpal femur laterally flattened. Female
first femur about 1.8-2.0 times carapace
length. Female calamistrum half the
length of metatarsus IV. Male first femur
with six ventroprolateral, three dorsopro-
lateral, one dorsoproximal, and three ret-
rolateral maerosetae (Fig. 39). Male first
tibia has one prolateral, three retrolateral,
one or two dorsal, and four ventral mae-
rosetae. Female abdomen oval, height
and width three-fourths length (Figs. 40,
42). Male abdomen cylindrical with di-
ameter half its length. A pair of stout tra-
cheal triuiks extends into cephalothorax.

Male Palpus. Palpal femur lacks ven-
tral tubercles and tarsus has an apparent-
ly reduced middle hematodocha. Median
apophysis a heavily sclerotized, elongate
projection with foot-shaped terminus that
appears to support the short, broad, blunt
embolus (Plate 3-A, B). Flat, curved radix
arises proximal to embolus and termi-
nates in two pointed prongs. Conductor
arises i^etween bases of median apophy-
sis and radix and extends between distal
pyortions of these sclerites.

Female Cenitalia. A posteroventral

hood of various widths (Figs. 32, 36, 43)
extends over dorsal atrium in which
openings are found (Figs. 33, 37, 44).
Each opening leads to a pair of appar-
ently blind spermathecae, the lateral pair
usually having thicker walls than the me-
dian pair (Figs. 34, 38, 45). Lateral to
spermathecal ducts is a pair of small (A.
albicans and A. mazolus, Figs. 34 and 38,
respectively) or large (A. aristus, Fig. 45)
accessory glands which appear to have
separate openings and to be homologous
with those of Waitkera, Folenecia, Hijp-
tiotes, and Siratoha.

Natural History and Distribution. The
three species in this genus are found
from southern Mexico through Central
America (Map 1). Nothing is known of
their web structure.

Key to Ariston Females
(Males are known only for A. mazolus)

1. Genitalia with a thin, anterior, overhanging
rim about three-fourths as wide as the
raised genital area; in ventral view posterior
genital margin rounded (Fig. 43); Panama
aristus

- Genitalia with a prominent median projec-
tion about one-fourth to one-third its width
and forming a hood; in ventral view poste-
rior genital margin indented (Figs. 32, 36);
southern Mexico to Honduras 2

2(1). Genital hood one-third the genital region's
width; in ventral \ iew this region's poste-
rior margin has a slight median depression
(Fig. 32) as does its dorsal margin when
viewed posteriorly (Fig. 33); clypeus height
one-fourth or less the AME diameter; south-
eastern Mexico and Honduras albicans

- Genital hood one-fourth the genital region's
width; in ventral view this region's poste-
rior margin has two lateral lobes separated
by a deep median depression (Fig. 36) and
in posterior view these lobes are separated
by a deep notch (Fig. 37); clypeus height
half or less the AME diameter; southwest-
ern Mexico niazoliis

Ariston albicans O. Pickard-Cambridge
Figures 32-35

.\ristoii (ilhicdii.s O. Pickard-Cambridge, 1896, Bio.
Cent.-Amer., Zool., Arachnida, 1: 161-224. Three
female syntypes from Teapa in the Mexican state
of Tabasco, collected by H. H. Smith, in the Brit-
ish Museum (Natural History), examined.

Revision of Uloboridae • Opell 481

Didf^nosis. Ariston albicans females
are distinguished from those of A. aristus
liy having a genital hood which is one-
third (Figs. 32, 33) rather than three-
fourths (Figs. 43, 44) the genital area's
width, by this hood forming a prominent
posteroventral projection rather than a
thin anterior margin of a broad atrium,
and by having an indented rather than
rounded posterior genital margin when
viewed ventrally. Accessory glands are
one-third (Fig. 34) rather than one-and-
one-half (Fig. 45) a spermathecal diame-
ter. Ariston albicans females differ from
those of A. niazohis by having a genital
hood which is one-third (Fig. 32) rather
than one-fourth (Fig. 36) the genital
area's width and by having shallow rather
than deep posterior and dorsal epigynal
indentations when viewed ventrally
(Figs. 32, 36) and posteriorly (Figs. 33,
37) respectively. Spermathecal ducts are
looped (Fig. 34) rather than straight
(Figs. 38, 45).

Description. Only females are known.
Total length 2.1 to 2.2 mm, carapace
length 0.7 mm, sternum length 0.5 mm.
Clypeus height one-tenth to one-fourth
AME diameter. Coloration similar to that
of A. aristus (Fig. 40). Carapace, sternum
and legs light tan. Dorsum of abdomen
white, with tan median cardiac region;
broad, posterior tan paraxial stripes and
a pair of tan spots just posterior to abdo-
men's center. From the center of the gen-
italia's ventral surface a narrow, flat, dor-
sally concave hood projects posteriorly
(Fig. 32). In ventral view (Fig. 32) genital
area's posterior edge slightly emarginate.
In posterior view (Fig. 33) genitalia's
broad, central region is concave and its
dorsal edge bordered by a thin, heavily
sclerotized, medially emarginate rim. A
genital opening is found at each ventro-
lateral margin of this dorsal rim and leads
to a short bursa from which a pair of
looped ducts originates. Each duct con-
nects to an apparently blind spermathe-
ca, the lateral one having thicker walls
than the median one (Fig. 34). Lateral to

the bursae is a pair of small, blind acces-
sory glands.

Distribution. Southeastern Mexico
and Honduras.

Ariston mazolus n. sp.

Figures 36-39, Plate 3-A, B

Types. Female holotype and three male paiatypes
from Miramar, Manzanillo in the Mexican state
of Colima, collected on 15 January 1943 In' F.
Bonet, in the American Museum of Natural His-
tory. The specific epithet is an arbitrary combi-
nation of letters.

Diagnosis. The only known males of
Ariston belong to this species and are
diagnosed in the generic treatment. Fe-
males are distinguished from those of A.
aristus by having a median genital pro-
jection which is one-fourth (Fig. 36) rath-
er than three-fourths (Fig. 43) the genital
area's width and by having an indented
rather than smooth posterior genital mar-
gin when viewed ventrally. Accessory
glands one-third (Fig. 38) rather than
one-and-one-half (Fig. 45) spermathecal
diameter. Ariston mazolus females differ
from those of A. albicans by having a
genital hood one-fourth (Fig. 36) rather
than one-third (Fig. 32) the genital area's
width, by having deep rather than shal-
low posterior and dorsal genital inden-
tations when viewed ventrally (Fig. 36)
and posteriorly (Fig. 37) respectively.
Spermathecal ducts straight (Fig. 38.)
rather than looped (Fig. 34).

Description. Male. Total length 1.4
mm, carapace length 0.6 to 0.7 mm, ster-
num length 0.4 mm. Carapace, sternum,
and legs tan. Al)domen with white dor-
sum and gray venter. Other features of
the male given in the genus description.

Female. Total length 2.0 mm, cara-
pace length 0.6 mm, sternum length 0.4
mm. Clypeus height half AME diameter.
Coloration similar to that of males. Gen-
italia's anteroventral region forms a
rounded ridge with an abrupt narrow me-
dian projection whose posterior surface
is concave (Fig. 37). In posterior view a
pair of lateral lobes separated by a deep

482 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

median notch is found dorsal to this pro-
jection (Fig. 37). Dorsal to these lobes is
a median lohe with a more heavily scler-
otized central region. A genital opening
located dorsolateral to each lateral lobe
leads via two ducts to a thick- walled lat-
eral and a thin-walled median sperma-
theca (Fig. 38). Near the bifurcation of
these ducts a small accessory gland is
found.

Distribution. Known only from the
type locality in southwestern Mexico.

Ariston aristus n. sp.
Figures 40-45

Types. Female liolotype and paratype from Barro
Colorado Island, Panama Canal Zone. Holotype
collected August 1939 by A. M. Chickering, para-
type 20-24 June by N. Banks, i)oth in the Mu-
seum of Comparative Zoology. The specific epi-
thet is an arbitrary combination of letters.

Diagnosis. Females are distinguished
by a broad, posteroventral genital atrium
bordered anteriorly by a narrow rim three-
fourths the genital area's width (Fig. 43).
In ventral view (Fig. 44) genitalia's pos-
terior edge gently rounded and not emar-
ginate as in other two species. Accessory
glands of A. aristus one-and-one-half the
spermathecal diameter (Fig. 45) rather
than one-third the diameter as in the other
two species (Figs. 34, 38).

Description. Only females are known.
Total length 2.3 to 2.5 mm, carapace
length 0.7 to 0.8 mm, sternum length 0.5
mm. Clypeus height half AME diameter.
Coloration (Fig. 40) similar to that of A.
aUncans. A thin, broad anterior rim on
the genital area's ventral surface borders
a broad, shallow atrium (Fig. 43). In pos-
terior view (Fig. 44) genital area's ventral
surface concave, its dorsal surface con-
vex, and its median dorsal margin with

two dorsal lobes separated by a shallow
depression. An opening located lateral to
the base of each lobe connects to a short
bursa giving rise to two straight ducts,
each leading to an apparently blind sper-
matheca (Fig. 45). A large accessory
gland lies ventral to each lateral sper-
matheca and opens externally via a sep-
arate duct.

Distribution. Known only from the
type locality in Panama.

Polenecia Lehtinen
Figures 46-54, Map 1

Stil)()ta [part]: — Simon, 1892, Histoire Naturelle des
Araign^es, 1(1): 1-256. Paris.

Polenecia Lehtinen, 1967, Ann. Zool. Fennici, 4:
199-468. Type species by original designation
and monotypy Uluboru.s productus Simon, 1873,
Mem. Soc. roy. sci. Liege, 2(5): 1-174. Seven ma-
ture and one immature female syntypes (No. AR-
150) from Corsica, in Museum National d'Histoire
Naturelle, Paris, examined. The genus name is
feminine.

Diagnosis. Polenecia and Sybota fe-
males are the only uloborids to have an
abdomen with a prominent posterior pro-
jection (Figs. 51, 102, 114, 116). Polene-
cia males and females lack PLE tubercles
(Fig. 50) like those found in male and fe-
male Sybota (Figs. 100, 103, 115) and are
distinguished by being the only ulobo-
rids to have a procurved anterior eye row
(Fig. 50). Males are characterized by a
palpus (Figs. 46, 47) with: 1. a thin cym-
bial projection and two short, marginal
cymbial macrosetae, 2. a broad conductor
with a long, thin medial projection
which, along with a thin proximal median
apophysis extension, serves as an embo-
lus guide, and 3. a lateral endite lobe
which bears stridulatory ridges. Polene-
cia females, unlike Sylyota females, have
a posteriorly directed, midventral geni-
tal extension (Fig. 52).

Figures 50-54. Polenecia producta (Simon). 50. Female carapace. 51. Lateral view of female abdomen. 52. Ventral view
of female genitalia. 53. Posterior view of female genitalia. 54. Dorsal view of cleared female genitalia. Figures 55-61.
Hyptiotes cavatus (Hentz). 55. Female carapace. 56. Lateral view of female abdomen. 57. Dorsal view of female, right,
first femur. 58. Retrolateral view of female left first patella showing position of lyriform organs. 59. Ventral view of
epigynum. 60. Posterior view of epigynum. 61 . Dorsal view of cleared epigynum. Figure 62. Hyptiotes paradoxus C. Koch,
dorsal view of cleared epigynum.

Revision of Uloboridae • Opell 483

484 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Description of Type Species. Female nerets and hooklung area. One male has
total length 3.8 mm, carapace length 1.2 a dark median, dorsal stripe. Male and
mm, sternum length 0.9 mm. Male total female carapace tan with white guanine
length 2.6 mm, carapace length 1.0 mm, deposits near thoracic depression and lat-
sternum length 0.8 mm. Carapace width eral margins of thoracic region. Sternum
and length eciual. Cephalic region rises has gray margins and brown central re-
from shallow thoracic depression and at gion. Legs without conspicuous mark-
level of ALEs has a width half maximum ings. Two stout tracheal trunks extend
carapace width. Although raised medial- inot cephalothorax.

ly, cephalic region has no conspicuous Male Palpus. Femur without ventral
eye tubercles (Fig. 50). Thoracic region tubercles and tarsus without middle he-
slopes abruptly to petiole. Anterior eye matodocha. Median apophysis' distal
row procurved. Posterior eye row re- lobe a flat plate and its proximal lobe a
curved such that a line across posterior long, flat spur which lies distal to the em-
margins of PMEs passes along anterior bolus and conforms to its curvature (Figs.
margins of PLEs. In dorsal view anterior 46, 47). Radix appears as a small lateral
extension of the clypeus equal to one plate near embolus base. A broad, flat
AME diameter and in anterior view conductor located at the proximal, medi-
height one to one-and-one-third AME di- an surface of the tarsus has two lobes: a
ameter. Median ocular area's anterior small lateral one and a long, thin, trough-
width half and its length two-thirds its shaped median lobe which, along with
posterior length. Female sternum width the median apophysis' proximal spur,
half and male sternum width 0.6 its serves as a guide for the embolus,
length. Prolateral surface of the female Female Genitalia. Genitalia a ventral
palpal femur concave. Female first femur mound with posteromedian lobe one-
1.5 and male first femur 1.6 times cara- third the width of raised genital area (Fig.
pace length. Female calamistrum 0.63 52). In posterior view (Fig. 53) genital
length of metatarsus IV. Male with weak- area's broad, weakly sclerotized dorso-
ly developed calamistrum two-thirds the median region bordered ventrally and
length of metatarsus IV. First male femur laterally by well sclerotized M-shaped
with two dorsal, four prolateral, and three edge of raised ventral region. An opening
retrolateral macrosetae (Fig. 49). First is found at each ventrolateral margin of
male tibia with two separate and four weakly sclerotized median region and
clustered ventral, one dorsal, two prolat- leads to a broad bursa giving rise to a
eral, and two retrolateral macrosetae. small, l)lind median bulb and a long.
Width and height of female abdomen coiled, anterior duct leading to a small,
two-thirds length. Prominent median tu- spherical, lateral spermatheca (Fig. 54).
bercle arises from posterior third of fe- Fertilization ducts may lead from these
male abdomen and extends posterodor- anterolateral spermathecae, but none is
sally for distance ec^ual to one-fourth to visible. Dorsal to spermathecal ducts is
one-third the abdomen's length (Fig. 51). a pair of large, apparently solid accessory
Male abdomen cylindrical, compressed glands whose ducts pass posteriorly, but
dorsoventrally and laterally constricted do not connect with the bursae. These
near center in a manner reminiscent of ducts probably have separate external
some casterine clubionids. A dorsome- openings as do those of Waitkera wait-
dian mound in the posterior third of kerensis.

male's abdomen bears a tuft of heavy se- Natural History and Distri]}ution. The

tae. Male and female abdomen white only described species, Poleneeia pro-

with dense covering of light setae and ducta, constructs a circular or semicir-

darkly pigmented around base of spin- cular vertical web which lacks capture

Revision of Uloboridae • Opell 485

spirals and has cribellate silk on the radii
and framework threads (Wiehle, 1931).
Eggsacs are attached to the twig which
forms a central, vertical support for the
web. This species is known from Corsica,
Spain, Portugal, Algeria, Tunisia, and
Syria (Map 1).

Hyptiotes Walckenaer

Figures 55-64, Plate 4-A-D; Map 1

Mithras [preoccupied] C. L. Koch, 1834, Arachni-
den. in Panzer, Faunae Insectorum Gemianiae
initia, 123: 9. Type species by monotypy Mithras
paradoxus C. L. Koch, 1834, ibid., disposition of
types unknown. Preoccupied by Mithras Hueb-
ner, 1818, Verz. bekannt. Schmett., 5: 79 — Lepi-
doptera.

Uptiotes Walckenaer, 1837, Histoire naturelle des
Insectes, Apteres, 1: 277-279. Type species by
monotypy Uptiotes anceps Walckenaer, 1837,
ibid. [= M. paradoxus C. L. Koch]. Disposition
of type specimens unknown.

Hyptiotes (emendation of Uptiotes Walckenaer,
1837, op. cit.): Erickson, 1845, Nomina systema-
tica Genervmi Arachnidanun, p. 14. in Agassiz,
1846, Nomen. Zool. Inde.x Univ. pp. 1-14.

CijUopodia Hentz, 1847, Boston J. Nat. Hist., 5: 466.
Type species by monotypy CijUopodia cavatus
Hentz, 1847. Female holotype from Alaliama, col-
lected in October by Hentz, specimen lost.

Note. The senior synonym Uptiotes
has not been used in primary literature
since 1845 and the emended name Hyp-
tiotes has gained universal acceptance.
Application is being made to The Inter-
national Commission on Zoological No-
menclature for suppression of the name
Uptiotes as an unused senior synonym
provisions of amended Article 79 of the
International Code of Zoological Nomen-
clature.

Dicifitiosis. Hijptiotes males and fe-
males are distinguished from those of all
other genera by having: 1. anterior half
of carapace abruptly narrowed to half the
width of the posterior half (Fig. 55), 2.
straight anterior eye row whose AMEs
are three to four diameters from cara-
pace's anterior rim, 3. median ocular area
whose anterior width is one-fourth its
posterior width, 4. carapace whose pos-
terior half is flat or depressed and in lat-

eral view slopes from petiole to PLEs,
and 5. length of female first femur less
than carapace length and male first femur
equal to carapace length.

Description. Total length of females
2.3 to 5.0 mm, of males 2.0 to 3.0 mm.
Carapace width equal to or slightly great-
er than length. Abdomen extends ante-
riorly over carapace, nearly reaching
level of posterior eye row in many
species. Posterior two-thirds of carapace
flat or medially depressed and sloping
upward from petiole to posterior eye row.
The PLEs on conspicuous lateral tuber-
cles and ALEs small and in some speci-
mens difficult to see (Fig. 55). Anterior
eye row straight and posterior eye row
recurved such that a line along posterior
margins of PMEs passes anterior to PLEs
by a distance equal to three-fourths to
one PLE diameter. Median ocular area's
anterior width one-fourth and its length
one-third its posterior width. In dorsal
view clypeus extends anteriorly a dis-
tance equal to four to five AME diame-
ters. Sternum width 0.5 to 0.6 its length.
Female palpal femur flattened. Female
first femur 0.7 to 0.9 carapace length.
Male first femur 0.9 to 1.1 carapace
length. Female calamistrum nine-tenths
length of metatarsus IV. Males have a
weak calamistrum four-fifths the length
of metatarsus IV. Male first femur with
two prolateral, three to seven retrolateral,
and three to four dorsal macrosetae (Fig.
63). Male first tibia with eight to twelve
prolateral, two to fifteen retrolateral,
three to four dorsal, and zero to seven
ventral macrosetae. Numerous stout se-
tae are also present on tibia I of most
males. Female abdomen two-thirds as
wide and two-fifth to three-fourths as
high as long (Fig. 56). Abdomen of many
species with four pairs of small, lateral
tubercles each bearing a tuft of flat setae.
Male abdomen usually more slender and
often lacking tubercles. Two stout tra-
cheal trunks extend into cephalothorax.

Male Palpus. Femur lacks proximal,
ventral tubercles and tarsus has no mid-

486

Bulletin Museum uf Cumparative Zoology, Vol. 148, No. 10

die hematodocha. Median apophysis
spur of American species forms a long,
thin, curved extension with a broad, flat-
tened tip (Plate 4-A-D). In Old World
species median apophysis is a short,
broad projection. Proximal portion of me-
dian apophysis flattened, expanded, and
covering most of tarsus' proximal lateral
surface. This flattened terminus lies in a
groove of the large, flat conductor and
with this sclerite serves as a guide for a
long, thin embolus which makes one-
and-one-half loops before terminating
near median apophysis spur (Fig. 64).
Between the grooved conductor terminus
and median apophysis spur's base is a
small, projecting sclerite that appears to
be a conductor branch.

Epigynum. Epigynum's posterior face
formed by a broad, sclerotized plate with
a narrow ventromedian extension con-
tinuing onto the epigynum's ventral sur-
face where it lies on a raised median
ridge and terminates in a prominent tu-
bercle (Figs. 59, 60). Epigynal openings
lateral to this extension's base, each lead-
ing via a broad bursa to a long, coiled
duct that presumably connects to the va-
gina (Figs. 61, 62). Among the coils of
each duct is a thick-walled accessory
gland (mistaken by Muma and Gertsch,
1964, for a spermatheca) whose long, thin
duct appears to open at the epigynum's
posterior margin independently of the
epigynal openings. In Old World species
(Fig. 62) the bursae are longer and the
accessory glands larger and more poste-
riorly situated than in American species
(Fig. 61).

Natural History and Distribution.
Members of this genus construct a verti-
cal, sectoral web consisting of four "ra-
dii," as discussed more fully under Nat-
ural History. This genus is represented

in North America, Europe, Japan, and,
according to Muma and Gertsch (1964),
India and Ceylon (Map 1).

Siratoba new genus

Figures 65-77, Plate 3-C, D; Map 1

Type. The type species of Siratoba is Ariston
referens Muma and Gertsch, 1964, Amer. Mus.
Novitates, 2196: 17. The genus name is an arbi-
trary combination of letters and is feminine.

Dia<ino.sis. Siratoba and Ariston are
the only American genera whose female
members have posterior genital hoods.
Female Siratoba have a total length of
3.0 mm or more and have a prominent
posterior plate in a genital atrium formed
i)y the ventral hood (Figs. 66, 76); where-
as, members of Ariston have a total
length of less than 2.6 mm and lack such
a plate (Figs. 33, 37, 44). Siratoba has
been collected only from the southwest-
ern United States and northern and cen-
tral Mexico. Ariston's range extends from
southern Mexico through Central Amer-
ica. Siratoba females are distinguished
by having: 1. PMEs separated by a dis-
tance no more than one-and-one-half
times that separating AMEs (Fig. 72), 2.
PME separation one-and-one-half times
that of the PME-PLE separation, 3. PME
and PLE mounds separate in anterior
view, and 4. abdomen with a middle peak
(Figs. 71, 74). The only known males be-
long to S. referena and are distinguished
by having a total length of about 2.7 mm
and a conspicuous, coiled radix with a
central groove in which the embolus lies
(Figs. 68, 69; Plate 3-C, D).

Description. Carapace width 0.80
length. Cephalic region rises slightly
from shallow thoracic depression and at
level of ALE is about 0.70 carapace
width. Both eye rows about equally re-
curved such that a line across posterior

Figures 63-64. Hyptlotes cavatus (Hentz). 63. Prolateral view of male, left, first femur, patella, and tibia. 64. Retrolateral
view of embolus (black), sperm reservoir, and fundus. Figures 65-74. Siratoba referena (Muma and Gertsch). 65. Ventral
view of epigynum. 66. Posterior view of epigynum. 67. Dorsal view of cleared epigynum. 68. Apical view of left male
palpus. 69. Retrolateral view of left male palpus. 70. Prolateral view of male, left, first femur, patella, and tibia. 71. Dorsal
view of female. 72. Female carapace. 73. Dorsal view of female rigfit first femur. 74. Lateral view of female abdomen.

Revision of Uloboridae • Opell 487

488 Bulletin Museui7i of Comparative Zoologij, Vol. 148, No. 10

margins of median eyes passes along an-
terior border of lateral eyes (Fig. 72).
Median ocular area about four-fifths as
wide anteriorly as posteriorly and its
length e(iual to its anterior width. In an-
terior view clypeus height of females
four-fifths to one AME diameter and in
males one-and-one-half AME diameter.
Sternum width 0.68 its length. Female
palpal lemur lateralh' flattened. Femur I
about 1.5 times carapace length. Female
calami strum half the length of metatarsus
IV. Femiu" I of males has six prolate ral,
three retrolateral, and one proximal dor-
sal macroseta (Fig. 70). Tibia I has three
prolateral, retrolateral, dorsal and ventral
macrosetae. Abdomen of male and fe-
male has a median central peak and its
width and height are each half its length
(Figs. 71, 74). Two stout tracheal trunks
extend into cephalothorax.

Male Palpus. Palpal femur bears a
proximal retrolateral lobe. Large, helical
radix with concave inner surface forming
a groove in which embolus lies (Figs. 68,
69; Plate 3-C, D). U-shaped conductor
lies along the radix's proximal surface
and its median lobe, along with part of
the radix, appears to rest in the trough-
shaped proximal median apophysis lobe.
The proximal lobe terminus in turn ap-
pears to rest in the grooved tip of the flat
distal median apophysis lobe.

Epigynum. A prominent, ventral hood
(Figs. 65, 75) extends posteriorly over a
dorsal atrium containing a median pos-
terior plate (Figs. 66, 76). An epigynal
opening is found at each ventrolateral
corner of this plate (Figs. 66, 76). A large,
thin-walled bursa leads from each open-
ing and connects to an oval, anterior sper-
matheca (Figs. 67, 77). Posterolateral to
the spermatheca is a smaller, spherical

accessory gland which appears to con-
nect both to the spermatheca's posterior
surface and to the exterior near the epig-
ynal opening. Connection between ac-
cessory glands and spermathecae is dif-
ficult to establish with certainty owing to
the small size of specimens.

Distribution. This genus is represent-
ed only in the southwestern United
States and northern and central Mexico
(Map 1). Nothing is known of its natural
history.

Key to Siratoba Females
(Males ot only S. referena are known)

1. Epigynal hood two-thirds as wide as the epig-
ynum (Fig. 65); in ventral view the epigynum's
posterior margin curved posteriorly, south-
western United States and northern Mexico
referena

- Epigynal hood one-third as wide as the epig-
ynum (Fig. 75); in ventral view the epigy-
num's posterior margin curved anteriorly, cen-
tral Mexico .sira

Siratoba referena (Muma and Gertsch)
new combination
Figures 65-74, Plate 3-C, D

Ariston rcjcrcns Muma and Gertsch, 1964, Amer.
Mus. Novitates, 2196; 17. Male holotype horn
Cochise Stronghold, Dragoon Mountains, Arizo-
na, collected 7 September 1950 by W. J. Gertsch,
female allotype from Douglas, Arizona, collected
27 August 1939 by R. H. Crandell, both in the
American Museum of Natural History, examined.

Didfinosis. The only known males of
Siratoba belong to this species. Females
are distinguished from those of S. sira by
having an epigynal hood which is two-
thirds (Fig. 65) rather than one-third (Fig.
75) the epigynal width and a posterior
epigynal margin which, in ventral view,
is curved posteriorly rather than an-
teriorly.

Figures 75-77. Siratoba sira n. sp. 75. Ventral view of epigynum. 76. Posterior view of epigynum. 77. Dorsal view of
cleared epigynum. Figures 78-85. Miagrammopes simus. 78. Dorsal view of female. 79. Lateral view of female cepha-
lothorax. 80. Dorsal view of male. 81. Female sternum, endites, and labium. 82. Male sternum, endites, and labium. 83.
Ventral view of epigynum. 84. Retrolateral view of female first left patella. 85. Retrolateral view of embolus (black), sperm
reservoir, and fundus. Figures 86-93. Miagrammopes latens Bryant. 86. Dorsal view of female. 87. Dorsal view of male.
88. Ventral view of epigynum. 89. Dorsal view of cleared epigynum. 90. Retrolateral view of left male palpus. 91 . Prolateral
view of left male palpus. 92. Female sternum, endites, and labium. 93. Male sternum, endites, and labium.

Revision of Uloboridae • Opell 489

490 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Description. Male. Total length 2.7 Description. Female. Total length

mm, carapace length 0.9 mm, sternum 2.9 mm, carapace length 0.9 mm, sternum

length 0.7 mm. Carapace and legs tan, length 0.6 mm. Carapace and legs light

sternum gra\'. Dorsum of abdomen white, tan, sternum light gray with dark border,

venter gray. Features of" the male palpus Dorsum of abdomen white, venter white

are given in genus description. with wide gray paraxial stripes. In ventral

Female. Total length 3.2 mm, cara- view (Fig. 75) epigynum's posteriorly di-
pace length 1.1 mm, sternum length 0.7 rected hood one-third its width and epig-
mm. Carapace and legs tan, sternum dark ynum's posterior margin curved anterior-
gray. Abdomen's dorsum white with ly. Hood with dorsal pit near its tip (Figs,
three pairs of posterior and one pair of 75, 76). In posterior view (Fig. 76) epig-
central gray spots (Fig. 71). Venter dark ynum's posterior margin concave and
gray with narrow longitudinal stripe ex- posterior plate's dorsal margin nearly
tending half the distance from epigastric straight. Sides of posterior plate nearly
furrow to cribellum. When viewed ven- parallel and epigynal openings located
trally (Fig. 65), epigynum has a poste- along their borders. Each opening leads
riorly directed hood two-thirds the epig- via a wide, thin-walled bursa to an oval
ynum's width and a posterior margin spermatheca. A small accessory gland ap-
which curves posteriorly. In posterior pears to connect to each spermatheca's
view (Fig. 66) epigynum's dorsal surface lateral margin by a short duct and to open
convex and posterior plate dorsally con- externally near the epigynal opening
cave, narrowing as it passes into hollow (Fig. 77).

formed by hood. Epigynal openings at Distribution. Central Mexico,
lateral margins of the posterior plate,

each leading to a large, thin-walled, bul- Miagrammopes O. Pickard-Cambridge
bous bursa which connects to a nearly Figures 78-97, Plate 5; Map 1
spherical spermatheca. A lateral, spheri-
cal accessory bulb opens lateral to each Miagrammopes O. Pickard-Cambridge, 1869, J.
epigynal opening and appears also to Linn. Soc. London (Zoo!.), 10: 400. Type species
connect via a convoluted duct to the sper- ^y ^""t"^' "^ fi""^* 'i^""^' >" publication M. thwaite-
matheca (Fig. 67). '" ^- Pi^^ard-Cambridge, 1869, ibid.

Distribution. Southwestern United Note. Octavius Pickard-Cambridge

States and northern Meixco. (1870) established the family Miagram-

mopidae for the genus Miagrammopes,

Siratoba sira n. sp. but Thorell (1873) considered Miagram-

Figures 75-77 mopinae as a uloborid subfemily. Lehti-

-r Filler ^ r ^ Hcu (1967) dividcs Miaarammopes (Mia-

liipes. Female holotvpe from entrance or cave at . , i r ii t^ \

Taninul in the Mexican state of San Luis Potosi, grammopmae) uito the toUowmg genera:

collected 29 March 1940 by W. Bridges, in the 1- MiagraniJllopes O. Pickard-Cam-

American Museum of Natural History. The spe- bridge, 1869, op. cit . Type species M

cific epithet is derived from the Greek term for thivuitesii O. Pickard-Cambridge, 1869,

op. cit. 2. Ran(i,unia Lehtinen, 1967, Ann.

Diagnosis. Males of this species are Zool. Fennici, 4: 199-468. Type species

unknown. Females are distinguished by original designation M. similis Kul-

from those S. refcrcna by having an epig- czynski, 1908, Ann. Mus. Nat. Hungary,

ynal hood which is one-third (Fig. 75) 6: 484. 3. Huanacauria Lehtinen, 1967,

rather than two-thirds (Fig. 65) the epig- op. cit. Type species by original desig-

ynum's width and a posterior epigynal nation M. bambusicola Simon, 1893,

margin which, when viewed ventrally, is Ann. Soc. ent. France, 61: 421-462. 4.

curved anteriorly rather than posteriorly. Mumaia Lehtinen, 1967, op. cit. Type

Revision of Uloboridae • Opell 491

106

108

Figure 94. Miagrammopes corticinus Simon, dorsal view of cleared epigynum. Figure 95. Miagrammopes bambusicola
Simon, dorsal view of cleared epigynum. Figures 96-97. Miagrammopes simus. 96. Dorsal view of cleared epigynum.
97. Dorsal view of female first right femur. Figures 98-108. Sybota abdominalis Nicolet. 98. Dorsal view of female. 99.
Dorsal view of female abdomen. 100. Female carapace. 101. Lateral view of female carapace. 102. Lateral view of female
abdomen. 103. Dorsal view of male. 104. Dorsal view of female first right femur. 105. Ventral view of epigynum. 106.
Posterior view of epigynum. 107. Ventral view of epigynum. 108. Posterior view of epigynum.

492 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

species by original designation Mia^nim- eter. The PME and PLE nearly equal in

mopes corticeus Simon, 1892, op. cit. size. The PLEs on distinct lateral tuber-

I have examined type specimens of cles. The PMEs may be from two to 20
Mia^rammopes corticeus, M. .similis, and times further from one another than from
M. hamhusicola as well as Neotropical, the PLEs (Figs. 78, 86). In addition to a
Australian and Oriental specimens of this shallow thoracic pit, a conspicuous
group and believe it to represent a mono- depression is located medially or pos-
phyletic assemblage. This is not to say teromedially to each PME. Sternum
that Lehtinen's division of this assem- with conspicuous intercoxal projections
blage is unwarranted. I have not under- and in many species divided by thin,
taken a comprehensive study of the mia- flexible, transverse regions into 3 plates:
grammopoid ulol)orids and for this reason one extending between the first two pairs
will attempt neither to characterize nor of coxae, a second l:)etween the third cox-
synonymize Lehtinen's genera. Rather, I ae, and a third between the fourth coxae,
will deal with this assemblage as the tra- Femur I 1.9 to 2.5 times as long as cara-
ditional or broader genus Mirtgrrtmmope.S' pace in females and 1.5 to 1.7 times in
sensu Into. My phylogenetic conclusions, males. Metatarsae I and IV laterally flat-
however, lead me to reject the subfamily tened and in females the latter has a row
ranking given to the group. of stout spines extending along most of

Di(i<i,nosis. Males and females of this its ventral surface and a dorsal calamis-

genus are distinguished from all other trum which is 0.6 to 0.7 its length. Femur

uloborid genera by lacking anterior eyes I of males lacks spines, but dorsal surface

(Figs. 78, 80, 86, 87), and by having a la- of tibia I has spines along its entire

biimi and endites which are two times as length. Abdomen of males and females

long as wide (Figs. 81, 82, 92, 93). The cylindrical to spindle-shaped, width 0.25

sternum of many species is very narrow to 0.40 length, usually attaining its max-

and divided into two or three plates by imum width and height near its center,

flexible transverse areas (Figs. 81, 92, 93). Transverse, common spiracular groove

Description. Total length of males 2.8 noticeably forward of cribellum, situated

to 3.5 mm, carapace length 1.0 to 1.2 mm, one-eighth to one-sixth distance from cri-

sternum length 0.5 to 0.6 mm. Total l^ellum to epigastric furrow. Two stout

length of females 4.0 to 6.2 mm, carapace tracheal trunks extend into cephalotho-

length 1.2 to 2.2 mm, sternum length 1.0 rax.

to 1.4 mm. Carapace width of both males Male Palpus. Femur lacks ventral, lat-
and females three-fourths to slightly eral tubercles. Long, narrow lobe on pa-
more than one times carapace length. In tella's dorsal surface extends above the
lateral view (Fig. 79) carapace nearly flat tarsal base (Plate 5- A, B; Fig. 90). He-
with first coxa extending from anterior matodocha absent. Median apophysis
surface. Anterior eyes absent. Posterior Indb and median apophysis spur modi-
median eyes separated from carapace's fied in both shape and orientation (Plate
anterior rim by a distance equal to 0.25 5). Conductor's proximal lobe serves as
to 0.36 the carapace length (Figs. 78, 80). an embolus guide, but its distal spur may
Posterior eye row either: 1. procurved be simple (Plate 5-A) or modified (Plate
such that a line across that anterior l:)or- 5-B-D).

der of the median eyes passes through Epigynum. Epigynum flat with open-

the lateral eyes or along their posterior ings at posterior. Each opening leads to

margin, 2. straight, or 3. recurved such a blind, anterolateral spermatheca whose

that a line across the median eyes' pos- posterior margin gives rise to either a fer-

terior margins passes anterior to the lat- tilization duct (Fig. 94) or a lateral bulb

eral eyes bv as much as one PLE diam- from which a fertilization duct arises

Revision of Uloboridae • Opell 493

(Figs. 89, 95). In M. simu.s (Fig. 96) each
epigynal opening leads first to a large,
oval bursa and then to a blind sperma-
theca.

Natural History and Distribution.
Members of this genus are known to pro-
duce either a single horizontal capture
thread with cribellar silk along its center
or a horizontal resting thread to which
one or several vertical or diagonal cribel-
lar capture threads are attached (Aker-
man, 1932; Lubin et al., 1978). The ge-
nus has a cosmotropical distribution.

Sybota Simon

Figures 98-119, Plate 6-A, B; Map 1

Sylvia [preoccupied] Nicolet, 1849, Arachnidos. /'n
Gay, Historia fisca y politica de Chile. Zoologia
3: 465. Type species Sylvid aI)dotninaJis Nicolet,
subsequent designation by Simon, 1892, Histoire
Naturelle des Araignees, 1(1): 216, Paris. Preoc-
cupied by Sylvia Scopoli 1769, Annus I: 154. —
Aves.

Sybota Simon, 1892, op. cit. New name (Feminine)
for homonym.

Diagnosis. Sybota and Polenecia fe-
males are the only uloborids to have a
posterior abdominal projection extending
beyond the spinnerets (Figs. 102, 110,
116). Sybota females and males have
prominent PLE tubercles (Figs. 100, 103,
115); whereas those of Polenecia do not
(Fig. 50). The anterior eye row of Sybota
males and females is recui-ved rather than
procui-ved as in Polenecia. Sybota males
are characterized by having: 1. no he-
matodocha (Plate 6-A, B), 2. a well-
developed conductor, and 3. a two- or
three-pronged median apophysis. The
posterior plate of Sybota females has a
ventrally directed median extension on
either side of which is a weakly sclero-
tized area (Figs. 105-108).

Description. Carapace width 0.7 to
1.0 its length. Female cephalic and tho-
racic regions each rise slightly from a
shallow thoracic depression and attain
about equal height (Fig. 101). Thoracic
region slopes steeply to petiole. Male
carapace nearly flat, rising slightly in

ocular area and sloping gradually from
thoracic pit to petiole. At ALEs cephalic
region of males and females is half as
wide as carapace (Figs. 101, 103). Ocular
area raised and conspicuously set off
from remainder of carapace (Figs. 100,
115). The PLEs on broad, prominent, lat-
erally directed tubercles. In S. mendoza
AMEs on an anterior tubercle (Fig. 115).
Both anterior and posterior eye rows are
recurved, posterior row such that a line
across the posterior margins of PMEs
passes along the anterior margins of
PLEs (Figs. 100, 115). In anterior view
female clypeus equal in height to AME
diameter and male clypeus 1.4 times
AME diameter. Male ALEs smaller than
AMEs. Median ocular area one-and-one-
half to two times as wide posteriorly as
anteriorly and about two-thirds as long as
its maximum width. Sternum width 0.6 to
0.7 its length. Female femur I 1.3 to 1.5
times as long as carapace; male femur I
1.5 to 1.7 times as long. Calamistrum half
the length of metatarsus IV. Female pal-
pal femur laterally compressed. Femur I
of males with three or four dorsal, five or
six prolateral, and three retrolateral
spines. Tibia I with four or five ventral,
two or three dorsal, seven proximal, and
two retrolateral spines. Female abdomen
half to three-fifths as wide and high as
long and posteriorly narrowed into a con-
ical projection which extends beyond
spinnerets for a distance equal to one-
fifth to one-third the abdomen's length
(Figs. 98, 102, 110, 114, 116). Male's ab-
domen oval, about half as wide as long,
dorsally flattened, and extended about
one-sixth its length beyond spinnerets
(Fig. 103). Four stout tracheal trunks ex-
tend into the cephalothorax.

Male Palpus. Femur without ventral
tubercles and tibia with a distal lobe
which extends beyond tarsal base. Long
dorsal spine present on patella and tibia.
Cymbium with two long spines on its dis-
tal median edge. Middle hematodocha
absent and palpal sclerites situated quite
distally (Plate 6-A, B). Median apophysis

494 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

terminal, flattened, and with three distal-
1\ directed processes. Emhohis curved
aroiHid median apophysis' base, coming
to He in the groove of conductor's l)asal
lol)e.

Epigynum. Epigynum's posterome-
dian margin formed by posterior phite
(Figs. 106, 113, 118) and, depending on
the shape of this phite, either rounded
(Fig. 112) or indented (Fig. 117) in ven-
tral view. Lateral to this plate on epigy-
num's ventral surface is a pair of level or
raised lighter areas, each with a low ridge
at its anterior border (Figs. 105, 106). An
epigynal opening is situated at each ven-
trolateral border of the posterior plate
and leads via a short duct to an anterior
bifurcation which connects with a blind
spermatheca and a posterior bifurcation
which gives rise to a fertilization duct
(Figs. 109, 119).

Natural History and Distribution. The
three known species in this genus appear
restricted to central and southern Chile
and the adjacent regions of Argentina
(xMap 1).

Key to Sybota
(Males of Sybota mendozae are not known.)

1. Males 4

- Females 2

2(1). Carapace width equal to length; AMEs not
on conspicuous tubercle (Figs. 98, 100);
posterior epigynal margin rounded slightly
(Figs. 105, 112); posterior plate pentagonal
or oval (Figs. 106, 113) 3

- Carapace width two-thirds length; AMEs
on prominent anterior tubercle (Figs. 114,
115); posterior epigynal margin indented
(Fig. 117); posterior plate diamond-shaped
(Fig. 118) mendozae

3(2). Posterior plate of epigyniun pentagonal, 1.5
times as broad as high and without a \'en-
tromedian depression (Figs. 105-108); car-
apace with white median line in cephalic
region and red pigment just anterior to tho-
racic depression (Fig. 98) ahdominalis

- Posterior plate oval, twice as broad as high
and with a slight ventromcdian depression
(Figs. 112, 113); carapace dark gray with no
white or red markings osoniis

4(1). Median apophysis of palpus with two prom-
inent projections and a small central spur at
the base of its broad lateral lobe (Plate 6-A);

dorsum of abdomen predominantly white

ahdominalis

- Median apophysis with three conspicuous
distal projections, the middle one clearly
set off (Plate 6-B); dorsum of abdomen dark
gray osornis

Sybota abdominalis (Nicolet)
Figures 98-109; Plate 6-A

Sylvia abdominalis Nicolet, 1849, Arachnidos. in
Gay, Historia fisca y poh'tica de Chile. Zoologia
3: 465-468. Female neotype from Pucatrihue in
the Chilean province of Osonoro, collected 12
April 1968 by L. Pena, in the Museum of Com-
parative Zoology.

Ulohoriis (d)dominaUs: — Simon, 1887, Ann. Soc.
ent. France, 7(6): 195.

Sybota ahdominalis: — .Simon, 1892, Histoire Na-
turelle des Araignees, 1(1): 1-256, Paris. New ge-
neric name for preoccupied Sylvia.

Note. Nicolet (1849) described five
species (Sylvia abdominalis, S. similis, S.
ater, S. rubig,inosa, and S. vittata) as be-
longing to his new genus Sylvia Ipreoc-
cupiedl. After examining Nicolet's spec-
imens, Simon (1892) concluded that they
were varieties of the same species and
selected S. abdominalis as the type
species for the replacement genus name
Sybota. As Lehtinen (1967) noted, the
Nicolet types no longer exist or, if they
do, cannot be located in the Museum Na-
tional d'Histoire Naturelle in Paris. It is
probably for this reason that all authors
prior to Lehtinen accepted Simon's in-
clusion of S. aJ)dominaIis and S. (Ulob-
orus) productus (Simon) in Sybota.
When Lehtinen created the genus Pole-
necia with P. productus as its type
species, he necessarily changed the
meaning of several studies dealing with
uloborid webs, e.g., Kaston (1964 and
1966) and Wiehle (1931). Because these
two genera play an important role in
studies of uloborid phylogeny and evo-
lution of web forms, it seems appropriate
to designate a neotype for S. abdomina-
lis. Additionally, description of two new
SyJ)ota species supports the need to
clearly associate this species name with
a type specimen. In the absence of a clear
definition of S. abdominalis I have as-

Revision of Uloboridae • Opell 495

sociated this name with the Chilean
species which appears more commonly
in collections, shows the more extensive
altitudinal and geographical distribution,
and has the greater color variation.

Diagnosis. Females of this species
are distinguished from those of S. men-
dozae by having a carapace width equal
to (Fig. 100) rather than two-thirds the
carapace length (Fig. 115), by having
only a slightly recurved anterior eye row,
and by having an epigynum whose pos-
terior margin is rounded (Figs. 105, 107)
rather than indented (Fig. 117), whose
posterior plate is pentagonal (Figs. 106,
' 108) rather than diamond-shaped (Fig.
118), and whose spermathecae are large
and elongate (Fig. 109). Sijhota ahdomi-
nalis females differ from those of S. osor-
nis by having a pentagonal posterior
plate (Figs. 106, 108) without a conspic-
uous ventromedian depression and by
having a carapace with a median white
line (Fig. 98) and a red pigment spot just
anterior to the thoracic depression.

Males of S. ahdominalis differ from
those of S. osornis by having a palpal
median apophysis with two (Plate 6- A)
rather than three (Plate 6-B) conspicuous
distal projections and by having white
rather (Fig. 103) than a dark gray abdom-
inal dorsum.

Description. Males. Total lengtli 3.6
to 4.0 mm, carapace length 1.3 to 1.4 mm,
sternum length 0.8 to 0.9 mm. Carapace
(Fig. 103) dark gray with tan lateral mar-
gins and a median white stripe extending
from thoracic depression to ocular region
where it expands. Sternum gray with a
central tan area. Abdomen (Fig. 103)
white with a dark gray posterior tip, three
pairs of dorsal gray spots, and lateral
transverse gray stripes. Median apophy-
sis of palpus with a long, thin projection
and a short, broad lobe with a small spur
on its mesal base (Plate 6- A).

Females. Total length 5.1 to 5.9 mm,
carapace length 1.4 to 1.6 mm, sternum
length 0.9 to 1.1 mm. Carapace (Fig. 98)
gray with a median white area extending

to ocular region. This white area is ex-
panded in the thoracic region of lighter
specimens and largely restricted to the
cephalic region of darker specimens. Red
pigment spot just anterior to thoracic
depression. Abdominal coloration vari-
able, ranging from completely white to a
gray venter and white dorsum (Fig. 98),
with or without a median dorsal gray
stripe; to a white venter and gray dorsum
with dorsal paraxial white stripes (Fig.
99). It is tempting to place specimens
with the latter coloration into a separate
species, but little other evidence sup-
ports this. Legs lack conspicuous mark-
ings and vary in color from tan to gray. In
ventral view (Figs. 105, 107) the epigy-
num's posterior margin is rounded and
lateral to the posterior plate's convex
ventral tip are two lightly sclerotized
areas, bordered laterally by dark, subsur-
face ducts. In posterior view (Figs. 106,
108) the posterior plate is pentagonal and
no more than 1.5 times as broad as high.
Ducts are short and looped, leading to
large, elongate spermathecae (Fig. 109).
Distribution. Collected from the
Chilean provinces of Osonio: Puyehue —
500 m, Pucatrihue— 200 m; Cautin: N.E.
Villarrica — 300 to 600 m; Llanquihue:
Chamiza — 0 to 100 m; Santiago: El
"Golf"; Concepcion: Bosque Ramunt-
cho— 0 to 100 m.

Sybota osornis n. sp.

Figures 110-113; Plate 6-B

Types. Female holotype and two male paratypes
from Purran<iue in the Chilean province of Osor-
no, eollected January-March 1955 l)y E. Reed, in
the American Museum of Natural History. The
specific epithet is a third declension noun in the
genitive case, derived from the province of the
type locality.

Diagnosis. Females of this species
are distinguished from those of S. men-
dozae by having a carapace width equal
to rather than two-thirds the carapace
length, by having only a slightly recurved
anterior eye row, and by having an epig-
ynum whose posterior margin is rounded

496 Bulletin Museum uf Comparative Zoulugij, Vol. 148, No. 10

(Fig. 112) rather than indented (Fig. 117), epigynum's posterior margin rounded.

whose posterior plate is oval (Fig. 113) Lateral to the posterior plate's concave

rather than diamond-shaped (Fig. 118), ventral tip are two lightly sclerotized

and whose spermathecae are large and areas, bordered laterally by dark, subsur-

elongate (Fig. 109). Syhota osoniis fe- face ducts. In posterior view (Fig. 113)

males differ from those of S. ahdominalis posterior plate oval and at least twice as

by having a broad oval posterior plate broad as high. Ducts short and looped,

with a conspicuous ventromedian leading to large, elongate spermathecae.

depression (Figs. 112, 113) and by having Distribution. Known only from the

a dark gray carapace with no color mark- type locality.

ings.

Males of S. osoniis differ from those of Sybota mendozae n. sp.

S. ahdominalis by having a palpal me- Figures 114-119

dian apophvsis with three (Plate 6-B) , , , , , ^ ,

.-,^K - 4^K t- tT>^ >- a \\ • Types. Female holotype and three female para-

rather than two Plate 6-A) conspicuous ' r 71 w e \a a ^ ^- 1

J . , 111 11 types from / km VV of Mendoza, Argentma, col-

distal projections and by having a dark Jected in "chapanal" at an altitude of 1200 m,

gray rather than a white abdominal dor- March-April 1958 hy B. Patterson. Holotype and

sum. two paratypes in the Museum of Comparative

Description. Male. Total length 3.8 Zoology, one paratype in the American Museum

. ^ 1 1 .1 1 F- . 1 .r. of Natural History. The specinc epithet is a nrst

to 4.1 mm, carapace length 1.5 to 1.6 mm, declension noun' in the genitive case, derived

sternum length 1.0 mm. Carapace darkly from the type locality.

mottled with gray except for light areas ^. ... 1 , t-

near the thoracic depression, in the ocu- D^^'^nosis. Males are not known. Fe-

lar area, and at the carapace margins dor- "]^^^^ '^'"^ distinguished from those of S.

sal to coxae III and IV. Sternum dark gray ^ahdominalis and S. osornis by having a

with a narrow median light stripe. Legs carapace whose width is two-thirds its

dark brown to gray, with no conspicuous ^^^^'^^ ^^'^^- ^l^' 115), by having AMEs

markings. Abdomen gray with small dor- °" ^ prominent tubercle, by having the

sal white spots and paraxial ventral 'Yiterior eye row more strongly recui-ved

stripes. Palpal median apophvsis with than the posterior, and by having an epig-

three distinct distal projecticms (Plate ynum whose posterior margin is indent-

g.g) ■ ed rather than rounded (Fig. 117), whose

Females. Total length 6.4 mm, cara- Posterior plate is diamond-shaped (Fig.

pace length 1.8 mm, sternum length 1.1 ^^^^ ^■'^*^^^^' **^'\" pentagonal or oval, and

mm. Carapace and sternum dark grav, whose spermathecae are small and spher-

sternum with a small tan center region. '^''^/ ''''*^^^'" ^'^'"^ ^'"S^^ '"^^ elongate (Fig.

Dorsum of abdomen dark gray with white 1 1 J).

anterior tip and two small, widely sepa- Description. Female. Total length 5.6

rated white spots in the anterior half, to 6.8 mm, carapace length 1.7 to 1.9 mm,

Venter with a median gray stripe and two sternum length 1.0 to 1.3 mm. Carapace

paraxial light stripes. A lateral white brown to light gray, lighter along anterior

stripe extends along each side of the ab- midline and darker in ocular region (Fig.

domen. Legs dark with no conspicuous 114). Sternum dark gray with brown an-

markings. In ventral view (Fig. 112) the terior median stripe. Dorsum of abdomen

Figure 109. Sybota abdcminalis Nicolet, dorsal view of cleared epigynum. Figures 110-113. Sybota osornis n. sp. 110.
Lateral view of female abdomen. 111. Retrolateral view of male left embolus (black), sperm reservoir, and fundus. 112.
Ventral view of epigynum. 113. Posterior view of epigynum. Figures 114-119. Sybota mendozae n. sp. 114. Dorsal view
of female. 115. Female carapace. 116. Lateral view of female abdomen. 117. Ventral view of epigynum. 118. Posterior
view of epigynum. 119. Dorsal view of cleared epigynum.

Revision of Uloboridae • Opell 497

^RES

115

113

118

498 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

all dark gray with slightly lighter paraxial
areas or white with dark gray anterior
median patch, gray transverse line in the
anterior third of its length, three pairs of
gray spots in the posterior half, and gray
posterior tip (Fig. 114). Venter of abdo-
men with a median gray longitudinal
stripe bordered by white patches or, in
dark specimens, all gray. Abdomen's lat-
eral surfaces all gray or white, mottled
with dark gray spots. Legs brown with a
dark gray longitudinal stripe on the dor-
sal surface of femur, patella, and tibia of
leg I. Femora, tibiae, and metatarsi of
legs II-IV each have a gray distal band.
In ventral view (Fig. 117) epigynum's
posterior margin indented with a pair of
slightly raised lateral lobes lateral to the
posterior plate's ventral tip. In posterior
view (Fig. 118) posterior plate is pentag-
onal. Ducts are long and highly convo-
luted, leading to small, spherical sper-
mathecae (Fig. 119).

Natural History and Distribution.
This species is known only from the type
locality in extreme western central Ar-
gentina. Darwin (1876) reported finding
several webs which "consisted of a
wedge-shaped segment" near Mendoza,
the type locality of this species. Morpho-
logical modifications characteristic of this
genus and species make its members
likely candidates for the production of
such apparently secondarily reduced
webs.

Orinomana Strand

Figures 120-128; Map 1

Oritwmus [preoccupied] Chamherlin, 1916, Bull.
Mus. Comp. Zoo!., 60(6): 206. Type species by
original desijination and monotypy O. lampru.s
Chamherlin, 1916, ihid. Preoccupied hy Orino-
mus Attems 1895, S.B. Akad. Wiss. Wien, Math.-
naturw. Ch, 104(1): 166.— Mvriapoda.

Oriuumanu Strand, 1934, Folia zool. hydrohioh,
6(2): 273. New name for Orinomus Chamherlin,
1916, preoccupied. The genus name is feminine.

Note. Mello-Leitao established the
genus Petrunkevitchia for P. venusta
Mello-Leitao, 1915, op. cit. To this genus
was added P. pusilla Mello-Leitao, 1917,
Arch. Esc. super, agric. medic, vet., 1(1):
3-19. Each species was described from
a single male specimen which apparently
no longer exists. Despite this, Lehtinen
(1967) treats this genus as a senior syn-
onym of Orinomana Strand, 1934 (nom.
nov. for the preoccupied Orinomus
Chamherlin, 1916, op. cit., containing
only the type species O. lamprus Cham-
herlin). However he considers P. pusilla
not to be congeneric with P. venusta, but
fails to assign it to another genus. On the
following page (258) Lehtinen places O.
lampra in the genus Philoponella. Ow-
ing to this confusion and inability to re-
solve the problem due to lack of speci-
mens of P. venusta and P. pusilla, I have
chosen to treat these two species and,
therefore, the genus Petrunkevitchia as
nomina dubia. The 1964 International
Code of Zoological Nomenclature de-
fines a nomen duhiuni as: "A name not
certainly applicable to any known taxon."
Mayr (1969) adds: ". . . owing to short-
comings in the original diagnosis or the
type material."

Diagnosis. Orinomana females, along
with those oiWaitkera and Sijbota, have
prominent PLE tubercles (Fig. 121). Ori-
nomana is distinguished from these gen-
era by having: 1. a peaked (Fig. 122) rath-
er than an oval (Fig. 24) or projecting
(Fig. 102) abdomen, and 2. AMEs which
are not on a slight, median mound and
which are one diameter removed from
the anterior carapace margin (Figs. 120,
121).

Figures 120-125. Orinomana bituberculata (Keyserling). 120. Dorsal view of female. 121. Female carapace. 122. Lateral
view of female abdomen. 123. Ventral view of epigynum. 124. Posterior view of epigynum. 125. Dorsal view of cleared
epigynum. Figures 126-128. Orinomana mana n. sp. 126. Ventral view of epigynum. 127. Posterior view of epigynum.
128. Dorsal view of cleared epigynum. Figures 129-130. Uloborus glomosus (Walckenaer). 129. Apical view of left male
palpus. 130. Dorsal view of cleared epigynum. Figure 131. Uloborus penicillatus Simon, dorsal view of male left first
femur.

Revision of Uloboridae • Opell 499

126

127

128

123

124

125

130

500 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Description. Carapace width and
length equal. Cephalic region rises
sharply from shallow thoracic depression
and thoracic region slopes abruptly to
petiole. The PLEs are on conspicuous
lateral tubercles and the entire ocular
area is on a distinct mound (Fig. 121). At
ALEs cephalic region 0.6 carapace width.
Both eye rows recurved so that a line
across the median eyes' posterior margin
passes through the anterior one-fourth of
the lateral eyes' diameter. Median ocular
area's anterior width three-fifths and its
length three-sevenths its posterior width.
In dorsal view clypeus extends forward
of AMEs by a distance equal to one AME
diameter. In anterior view clypeus height
four-fifths an AME diameter. Sternum 0.6
as wide as long. Female palpal femur flat-
tened. First femur 1.4 times as long as
carapace. Calamistrum 0.7 times as long
as metatarsus IV. Abdomen height and
length nearly equal, width two-thirds to
three-fourths length (Figs. 120, 122).
Dorsum with a pair of large anterior tu-
bercles and two pairs of small, parame-
dial tubercles posterior to these. Two
stout tracheal trunks extend into cepha-
lothorax.

Epigynum. Epigynum with a pos-
teromedian notch or indentation and a
pair of posterolateral depressions (Figs.
123, 124, 126, 127). An opening is found
in each depression and the straight duct
leading from it divides to connect to
blind anterolateral spermatheca and
either a long fertilization duct or a pos-
terior spermatheca with a posteromedian
fetilization duct (Figs. 125, 128).-

Natural History and Distribution.
Nothing is known of the natural history
of this genus. Its members are found in
the high Andes (3000 to 4000 m) from
southern Ecuador to northern Chile.

Key to Females of Orinomana
(No males of this genus are known)

1. Sternum uniform brown; posterior epigynal
margin with a sharp, V-shaped median noteh
(Fig. 123); abdomen white (Fig. 120)
hituherculata

- Sternum lirown with a light median stripe;
posterior epigynal margin with a rounded me-
dian indentation (Fig. 126); abdomen darkly
colored inana

Orinomana bituberculata (Keyserling),
new combination
Figures 120-125

Ulohorus hituherculatiis Keyserling, 1882, Verh.
zt)()l.-bot. Ges. Wien, 31: 282. Two identified fe-
males from Peru in a vial labeled "type" in Brit-
ish Museum (Natinal History), examined. Two
identified females in vial labeled "Lima, Peru,
leg. K. Yebki? Y. Sublemaa? detm. E. Keyserling,
177," in Polska Akademia Nauk Instytut Zoolo-
giczny, Warsaw where Keyserling (1882 op. cit.)
noted specimens to be, examined.

Orinomus himpriis Chamberlin, 1916, Bull. Mus.
Comp. Zool., 60(6): 207. Female holotype from
Urubamba (3160 m), Peru, collected 1919, in the
Museum of Comparative Zoology, examined,
NEW SYNONYMY.

Orinomana lampra: — Strand, 1934, Folia zool. hy-
drobiol., 6(2): 273.

Diafinosis. Orinomana hituherculata
females are distinguished from those of
O. niana by having a brown, unmarked
sternum, a white abdomen (Fig. 120), a
sharply notched posterior epigynal mar-
gin (Fig. 123), and two pairs of oval sper-
mathecae (Fig. 125).

Description. Only females are known.
Total length 4.8 mm, carapace length 1.2
mm, sternum length 0.8 mm. Carapace
mottled brown with thin, light median
stripe, a light ocular area with black en-
circled eyes, and a thin, light lateral mar-
gin (Fig. 120). Leg I tan mottled with
gray; femur and tibia with faint proximal
and median light rings; proximal two-
thirds of metatarsus nearly white. Abdo-
men white with faint gray patches lateral
to each dorsal tubercle (Fig. 120).

Epigynum. In ventral view (Fig. 123)
posterior epigynal margin with a deep
median notch and two broad, lateral
depressions. Posterior epigynal surface
(Fig. 124) nearly flat except for ventro-
median notch. Two pairs of nearly equal-
sized oval spermathecae (Fig. 125).

Di.strihution. This species is known
only from the type locality in central Peru
and from a female collected at Cerro Ti-

Revision of Uloboridae • Opell 501

najillas (3100 m) in the southern Ecua-
doran province of Azuay on 18 to 21
March 1965 by L. Pena.

Orinomana mana n. sp.
Figures 126-128

Type. Female holotype from Quismaa (ca. 4000 m)
in the Chilean province of Tarapace, collected 5
June 1968 by L. Pena, in the Museum of Com-
parative Zoology. The specific name is an arbi-
trary combination of letters.

Diagnosis. Orinomana mana females
are distinguished from those of O. ])itu-
berculata by having a Hght median ster-
nal stripe, a darkly marked abdomen, a
rounded posteromedian epigynal inden-
tation (Fig. 126), and a single pair of sper-
mathecae (Fig. 128).

Description. Only females are known.
Total length 3.8 mm, carapace length 1.1
mm, sternum length 0.8 mm. Carapace
coloration similar to O. hituherculata but
the median stripe is broader and the lat-
eral ocular area is brown not white. Car-
apace gray with a light median stripe.
Leg I light brown with a light median
femoral and a proximal tibial ring and a
brown metatarsus. Dorsum of abdomen
white overlain by dense black mottling
to form a dark, median stripe and a dark
chevron between each pair of light, dor-
sal tubercles. Venter of abdomen black
with a thin median and a wide paraxial
light stripe.

Epigynum. In ventral view (Fig. 126)
posterior epigynal margin with a rounded
median indentation containing two small
central projections. In posterior view
(Fig. 127) these projections appear dor-
sally directed. A single pair of sperma-
thecae is present (Fig. 128).

Distribution. Known only from the
type locality in northern Chile.

Uloborus Latreille

Figures 129-156, Plate 7-A; Map 2

Uloborus Latreille, 1806, Genera Crustaceorum et
Insectorum, Araneides, 1: 109, Paris. Type
species by monotypy U. walckcuaerius Latreille,
1806, i])id. Tvpe specimens no longer exist.

PhiUijra Hentz, 1850, Boston J. Natin-. Hist. Soc, 6:
24. Type species l)y virtue of first listing in pub-
lication P. mammeata Hentz, 18.50, ibid. The ho-
lotype of P. mammeata does not exist, but this
species has long been recognized as a synonym
of Uloborus glomosus (Walckenaer).

Veleda Blackwall, 1859, Ann. Mag. Natur. Hist., ser.
3, .3: 95. Type species by monotypy V. lineata
Blackwall, 1859, ibid. The type species is a syn-
onym of U. walckenaeriiis.

Philoponus Thorell, 1887, Ann. Mus. Civ. stor. nat.
Genova, ser. 2, 5: 127. Type species by monotypy
Philoponus pteropus Thorell, 1887, i])id.

Diagnosis. Uloborus males are distin-
guished by having: 1. a pear-shaped car-
apace with conspicuously narrowed ce-
phalic region which is two-fifths the
maximum carapace width (Figs. 133, 141,
150), 2. a posterior eye row recurved such
that a line across posterior margin of
PMEs passes anterior to the PLEs by half
a PLE diameter, 3. clypeus height e(iual
to half an AIVIE diameter, 4. a conductor
with only a proximal lobe (Plate 7-A, Fig.
129) and 5. numerous trichobothria on
the first pairs of legs (Fig. 131). Females
are similar to those oi Octonarius, having
slender posterolateral epigynal lol)es
whose bases are contiguous (Figs. 137,
145). Unlike Octonarius, they: 1. lack
conspicuous posterodorsal atria (Figs.
138, 146), 2. have a cephalic region which
at the ALEs is only half as wide as the
carapace (Fig. 132), 3. have more con-
spicuously recurved eye rows, and 4.
have a tuft of setae on distal region of the
first tibia (Figs. 136, 153).

Description. Carapace width equal to
0.85 length. Cephalic and thoracic re-
gions of females level, thoracic depres-
sion a shallow pit. Cephalic region of
males curved slightly downward from
thoracic depression. Both eye rows re-
cui-ved. A line across AMEs' posterior
margins passes anterior to the ALEs by
a distance equal to one-fourth an ALE
diameter in females and one ALE diam-
eter in males. A line across PME's pos-
terior margins passes anterior to the
PLEs by a distance equal to one-fourth
a PLE diameter in females and half a
PLE diameter in males. Median ocular

502 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

area is square. In anterior view clypeiis terior margin a fertilization duct leads
height equal to two-fifths to half an AME (Figs. 130, 139, 147, 156).
diameter. Sternum width 0.72 its length. Natural History and Distribution. As
Female palpal femur laterally com- far as is known, all members of this cos-
pressed. Length of first femur 1.4 to 1.5 mopolitan genus spin horizontal orb-
that of carapace. Female calamistrum half webs.

as long as metatarsus IV. Female first tib- Note. Uloborus festivus Mello-Leitao

ia with a distal tuft of setae forming a con- female holotype is an immature speci-

spicuous brush. Male first femur with two men, U. orsinus Mello-Leitao female ho-

prolateral, two retrolateral, and two dor- lotype has no abdomen, and U. peruva-

sal spines. Male first tibia with ten pro- nu.s Keyserling female holotype has been

lateral, six retrolateral, and 12 dorsal dried and its genitalia cannot be studied,

spines. Female abdomen three-fourths as These three species must be treated as

high and half as wide as long, usually nomina duhia.

with one pair of dorsal tubercles in the Confusion over the large number of

anterior third of its length. Male's abdo- Uloborus species names in the literature

men half as wide and high as long and (most appearing only in original species

without tubercles. Tracheal system with descriptions and many belonging to what

either two or four stout tracheal trunks are now recognized as other genera)

extending into cephalothorax. seems to have prevented many arachnol-

Male Palpus. Femur with a pair of ogists from ascribing any name at all to

proximal, ventral tubercles and tarsus members of this genus. Although the

with a prominent middle hematodocha three Uloborus species redescribed in

(Plate 7-A). Median apophysis bulb well this study are clearly distinct from one

developed and bears a terminal median another, each includes considerable col-

apophysis spur. Conductor arises from or variability and may actually represent

median apophysis bulb's base and lacks several sibling species which I am un-

the distal spur present in Philoponella able to distinguish at this time. This seems

males. In U. sef^rc^atus and U. campes- most likely in U. aegrotus and \edst\ike\y

tratus the conductor is as long as broad in U. cinereus. With synonyms accounted

(Figs. 143, 151); whereas, in U. penicil- for and the following three species delin-

latus it is much longer than broad (Fig. eated, names can be consistently ascribed

135). to specimens and, as more information

Epigynum. Two weakly sclerotized becomes available, the validity of the

posterolateral lobes extend from epigy- species described here more rigorously

num's posterior margin (Figs. 137, 154). tested.

A small sclerite is usually found at the tip

of each lobe and a copulatory opening is Key to Uloborus Males

found dorsal or dorsolateral to the base of , ^^ i . i i ^ ^ ^ c ^- ^

, , , r-' 1 ■ 1 1 • 1 1- Conductor lobe at least tour tunes as long

each lobe. Each openmg leads via a duct .^^ ^^,i^ (pj^, 135). carapace usually with

to an oval spermatheca from whose pos- three light stripes (Fig. 133)___- pcnicilhitus

Figures 132-139. Uloborus penicillatus Simon. 132. Dorsal view of female. 133. Dorsal view of male. 134. Ventral view
of male left palpal femur. 135. Apical view of male left palpus. 136. Retrolateral view of female left first leg. 137. Ventral
view of epigynum. 138. Posterior view of epigynum. 139. Dorsal view of cleared epigynum. Figures 140-147. Uloborus
segregatus Gertsch. 140. Dorsal view of female. 141. Dorsal view of male. 142. Dorsal view of male. 143. Apical view of
male left palpus. 144. Ventral view of male left palpal femur. 145. Ventral view of epigynum. 146. Posterior view of
epigynum. 147. Dorsal view of cleared epigynum. Figures 148-156. Uloborus campestratus Simon. 148. Dorsal view of
female. 149. Dorsal view of female abdomen. 150. Dorsal view of male. 151. Apical view of male palpus. 152. Ventral view
of male left palpal femur. 153. Retrolateral view of male left first leg. 154. Ventral view of epigynum. 155. Posterior view
of epigynum. 156. Dorsal view of cleared epigynum.

Revision of Uloboridae • Opell 503

504 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

- Conductor lobe as long as wide (Fij^s. 143,
151); carapace with a single median stripe
(Fig. 141) or, if with three stripes a central
guanine spot is present (Fig. 150) 2

2(1). Carapace with a central guanine spot (Fig.
150); median stripe broad; median apoph-
ysis bulb crescent-shaped, its prolateral

notch t\\ ice as broad as deep (Fig. 151)

campestratus

- Carapace without a central white spot (Fig.
141, 142); median stripe narrow; median
apoph\'sis bid!) semicircular, its prolateral

notch as broad as deep (Fig. 143)

segregatus

Key to Uloborus Females

1. Epig>nal lobes separateK- connected to
epigviial moiuid with their bases contig-
uous, forming a deep median "V" (Fig.
145); each lobe usually has a large, heavily
sclerotized cap segregatus

- Epigynal lobes have a common connection
to epigNiial mound with their bases fonning
a shallow median "U" (Fig. 137) or their
bases separated from one another (Fig.
154); lobes with common base lack a large
distal cap 2

2(1). Carapace with a large, central guanine spot
and often with a broad, central light median
stripe (Fig. 148); abdomen usually with four
pairs of dorsal humps, each hiuiip beset
with a tuft of thick white setae (Figs. 148,
149); first tibia with a dark, distal band
which in a few lighter specimens is very
faint (Fig. 153) campestratus

- Carapace without a white spot, lighter spec-
imens have a narrow median white stripe
and two paraxial white stripes (Fig. 132),
darker specimens have only a central white
stripe or a white posteromedian wedge; ab-
domen with only a single pair of anterior
humps (Fig. 132); tibia I unmarked (Fig.
136) or, in dark specimens with a very nar-
row proximal i)and penicillatus

Uloborus penicillatus Simon,
new combination
Figures 131-139; Map 2

uloborus penicillatus Simon, 1891, Proc. Zool. Soc.
London, 1891: 5.54. Three female, one male, and
immature syntypes (in three vials) from the Brit-
ish West Indies island of St. Vincent, collected
by H. H. Smith, in the British Museum (Natural
History), examined.

Uloborus aegrotus Simon, 1893, Ann. Soc. ent.
France, 61: 424. Three female and three imma-
ture syntypes from Caracas and San Esteban,
Venezuela, collected 27 December 1887 to 20
January 1888 and 29 February to 27 March 1888,
respectively by E. Simon, in Museum National
d'Histoire Naturelle, Paris, examined. NEW
SYNONYMY.

Uloborus Duiniculatus Simon, 1893, Ann. Soc. ent.
France, 62: 299. Two female and one immature
s\nt\pes from San Paulo de Olivenca (de Mathan)
in tiic Brazilian state of Amazonas, AR 45, in the
Museum National d'Histoire Naturelle, Paris, ex-
amined, NEW SYNONYMY.

Uloborus bucki Mello-Leitao, 1943, Arquivos do
Museu Nacional, Rio de Janeiro, 37: 157. Female
holotype from Porto Algere, Rio Grande do Sul,
Brazil, collected by P. Buck, in Museu Nacional,
Rio de Janeiro (No. 41. 720), examined, NEW
SYNONYMY.

Uloborus plurnipedatus Roewer, 1951, Abh. Nat.
Ver. Bremen, 32(2): 154. New name for U. plu-
mipes Mello-Leitao, 1947, Boletin Museu Na-
cional, Rio de Janeiro, 80: 6. Preoccupied by U.
plumipes Lucas, 1846, Explor. Sci. Algerie, Zool.
1. Araneides, p. 252. Two female, one immature,
and two male syntypes from Garmo do Rio Claro
in the Brazilian state of Minas Gerais, collected
by J. C. Carvalho, in the Museu Nacional, Rio de
Janeiro, examined, NEW SYNONYMY.

Diagnosis. In many respects this
species is similar to U. gjomosus from
North America (Muma and Gertsch,
1964). Males are distinguished hy having
a conductor which is about five times as
long as wide (Fig. 135) rather than three
times as long as wide (Fig. 129) and by
having a small median apophysis bulb
with a deep prolateral notch (Fig. 135)
rather than a large median apophysis
bulb with a shallow notch (Fig. 129).
Males are more easily distinguished from
those of U. .seff regains and U. campestra-
tus by their long conductor (Fig. 135) and
by the presence of small, less erect setae
on the shaft of the palpal femur (Fig.
134). Uloborus penicillatus females are
more difficult to distinguish from those of
U. gloniosus. The former usually lacks a
proximal light tibial ring in all but dark
specimens; whereas, most specimens of
the latter have a conspicuous, though nar-
row, proximal light tibial ring. Epigynal
lobes of U. penicillatus unite medially
before joining the epigynal (Fig. 137)
mound while those of most U. glomosus
unite at the epigynal mound. Uloborus
penicillatus females also differ from
those of U. segregatus and U. campestra-
tus, whose epigynal lobes unite at the
epigynal mound (Fig. 145) or are separate
at their attachment with the epigynal

Revision of Uloboridae • Opell

505

\ "^

^^ V^ - N.

g-^i^-X

^;p5\^/\j

V-l ^K^r:''^'— >-i-ta

yp / 'if ■ ^'^N ^W

^4— -v*v^~~"*^

^n5*s "■-- Tv,

p*^' /"' >.':•. ^iS ^W

^^*'"^-i ^S«fs

ffT"

^%^c

) ..---.

^:H V:::i J

r^^^i ■

S*^ >^- ^i. ■ y-^

T 7'"; •/

i^-w

•fv'^^-- ^ ^

/? s

4'-^^ - '^^^

y n /

J* ^ K±^^

1 l-v \J

oj; j?"^

u ,j-^

r^j-^ j^

^*iA. "^

• Uloborus penicillatus

**^ "''

Uloborus

campestratus

o Uloborus segregafus

Zosis geniculatus

Map 2. Distribution of Uloborus campestratus, U. penicillatus, U. segregatus, and ZosIs geniculatus.

mound (Fig. 154). Uloborus penicillatus
females lack the carapace guanine spot
(Fig. 148), dark, distal tibial ring (Fig.
153), and four pairs of abdominal tuber-
cles (Figs. 148, 149) found in U. campes-
tratus. Epigynal lobes of U. penicillatus
(Fig. 137) lack well sclerotized tips pres-
ent in most U. segregatus specimens
(Fig. 145).

Description. Female total length 4.4
to 7.2 mm, carapace length 1.4 to 1.7 mm,
sternum length 0.8 to 1.2 mm. Male total
length 3.0 to 4.2 mm, carapace length 1.1
to 1.3 mm, sternum length 0.6 to 0.9 mm.
Female carapace tan to dark gray, lighter
specimens with a light narrow median
stripe and two light paraxial stripes (Fig.
132), darker specimens with only a me-
dian stripe. Male carapace tan with light
median and paraxial stripes (Fig. 133).
Female abdomen tan (Fig. 132) to dark
gray. Male abdomen usually tan (Fig.
133), in a few specimens dusty gray. Two
stout tracheal trunks extend into cepha-
lothorax.

Male Palpus. Palpal femur's large ret-
rolateral tubercle (Fig. 134) directed

more perpendicular to femur's axis than
in U. segregatus or U. campestratus
(Figs. 144, 152). All setae but the one
arising from the tubercle's tip smaller
and less erect than in the other two
species. Median apophysis bulb small
with a deep prolate ral notch (Fig. 135).
Conductor at least five times as long as
broad.

Epigynum. Epigynal lobes slender
uniting at their bases to form a "U" which
is often lighter than the remainder of the
epigynum to which the lobes' common
base attaches (Figs. 137, 138). Sperma-
thecae oval, ducts long and convoluted
(Fig. 139).

Natural History and Distribution. As
far as is known members of this species
construct tubular eggsacs with lengths at
least ten times their diameters. This
species is known from southern Mexico
through Central America to Ecuador, Par-
aguay, and southeastern Brazil (Map 2).

Uloborus segregatus Gertsch
Figures 140-147; Map 2

uloborus segregatus Gertsch, 1936, Amer. Mus.

506 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Novitates, 852: 4. Male li()l()t>i)t' and two male
paratypes from EdinhuiK, Texas, collected 16
September 1935 by S. Mulaik. Two female para-
types from fi\e miles SE of Welasco, Texas, col-
lected 21 July 1935 by S. Mulaik. In the American
Museum of Natural History, examined. Muma
and Gertsch, 1964, Amer. Mus. Novitates,
2196: 26.

Diagnosi.s. As Muma and Gertsch
(1964) note, males of this species are dis-
tinguished from those of U. divcrsus l)y
havinsj; a palpal femur which is 2.5 rather
than 4.5 times as long as broad. Females
lack a dark, distal tibial ring present in U.
diversus. Ulohorus segregatus males dif-
fer from those of U. penicillatus by hav-
ing a short conductor (Fig. 143), erect se-
tae on the palpal femoral shaft (Fig. 144),
and a single longitudinal stripe on the
carapace (Figs. 141, 142). They differ
from U. cam pest ratus males by lacking
a central white spot on the carapace's
center and by having a larger retrolateral
palpal femoral tubercle. Ulohorus segre-
gatus females are distinguished from
those of U. penicillatus by having epig-
ynal lobes which unite to form a "V" rath-
er than a "U," and have sclerotized tips
(Figs. 145, 146) and from those of U. cam-
pestratus by lacking a central guanine
spot at the carapace's center and a dark,
distal ring on the first tibia.

Description. Female total length 2.8
to 4.2 mm, carapace length 0.8 to 1.4 mm,
sternum length 0.6 to 0.9 mm. Male total
length 2.4 to 3.0 mm, carapace length 0.8
to 1.0 mm, sternum length 0.5 to 0.6 mm.
Female and male carapace gray with a
light median stripe. Abdomen of most fe-
males and darker males light dorsal ly and
gray laterally with a complete or broken
median gray stripe (Figs. 140, 141).
Lighter males with a tan abdomen. Four
stout tracheal trunks extend into the
cephalothorax.

iMale Palpus. Femoral shaft with six
large, erect setae (Fig. 144). Median
apophysis with a deep prolateral notch
(Fig. 143). Conductor length and width
equal.

Epigynum. Epigynal lobes unite sep-

arately to epigynal mound with their bas-
es contiguous (Figs. 145, 146). Each lobe
usually has a conspicuous sclerotized
cap. Spermathecae round, ducts short
(Fig. 147).

Natural History and Distribution.
Vials of several specimens contained
stellate eggcases similar to that shown in
Fig. 233. This species is found from
southeastern Texas through Colombia
(Map 2).

Uloborus campestratus Simon,
new combination
Figures 148-156; Map 2

ulohorus campestratus Simon, 1893, Ann. Soc. ent.
France, 61: 424. Nine female and three male (see
Note) syntypes from La Guiara, Caracas, and San
Esteban, Venezuela collected by E. Simon from
27 December 1887 to 27 March 1888 in Mu-
seum National d'Histoire Naturelle, Paris, ex-
amined.

Uloborus cinercus O. Pickard-Cambridge, 1898,
Biol. Cent. -Amer., Zool., Arachnida. Araneida, 1:
265. Two female syntypes from Teapa in the
Mexican state of Tal)asco, collected by H. H.
Smith, in British Museum (Natural History), ex-
amined, NEW SYNONYMY. Muma and Gertsch,
1964, Amer. Mus. Novitates, 2196: 28.

Ulohorus spernax O. Pickard-Cambridge, 1898, op.
fit. Female holotype from Teapa in the Mexican
state of Tabasco, collected l)y H. H. Smith, in the
British Museum (Natural History), examined,
NEW SYNONYMY.

Note. Type material of U. campestra-
tus contained nine females conspecific
with U. cinereus and V. spernax plus
three males conspecific with Philoponel-
la fasciata. Simon describes females be-
fore males, and for this reason, I assert
my authority as first revisor in establish-
ing the synonymy listed above.

Diagnosis. Males and females are dis-
tinguished from those of other species by
the presence of a large, white guanine
spot at the carapace's center (Figs. 148,
150). Ulohorus campestratus females are
the only members of this genus to have
loin- pairs of dorsal abdominal tubercles
(Figs. 148, 149) and a dark ring on the
distal half of the first tibia (Fig. 153).

Description. Female total length 2.4

Revision of Uloboridae • Opell 507

to 4.0 mm, carapace length 0.8 to 1.3 mm,
sternum length 0.5 to 0.8 mm. Male total
length 2.0 to 2.4 mm, carapace length 0.9
mm, sternum 0.4 mm. Female carapace
light tan to gray (Fig. 148), darker speci-
mens with a broad, median light stripe.
Male carapace of darker specimens with
a broad median and two posterior parax-
ial stripes (Fig. 150). Female abdomen
light tan (Fig. 149) to light gray, darker
specimens with a median gray stripe and
a pair of lobed, paraxial white stripes ex-
tending laterally to abdominal tubercles
(Fig. 148). Male abdomen tan, without
dorsal humps. Four stout tracheal trunks
extend into cephalothorax.

Male Palpus. Palpal femur's retrolat-
eral tubercle (Fig. 152) smaller than that
of U. penicillatus and U. seg,reg,atus
(Figs. 134, 144). Median apophysis bulb
small, with a broad, shallow prolateral
notch (Fig. 151). Conductor length and
width equal.

Epigynum. Epigynal lobes broad and
medially, widely separated at bases
(Figs. 154, 155). Spermathecae oval with
short ducts (Fig. 156).

Distribution. Florida and Gulf Coast
states through Central America to Vene-
zuela (Map 2).

Purumitra Lehtinen

Figures 157-161; Map 1

Purumitra Lehtinen, 1967, Ann. Zool. Fennici, 4:
261. Type species by original designation and
monotypy Uloborus ^rammicus Simon, 1893,
Ann. Soc. ent. Fiance, 62: 68. Male holotype from
Manila, Philippine Islands, in Museum National
d'Histoire Natinelle, examined. The genus name
is feminine.

Note. Three female and two male
specimens of P. ^^rammicus, collected
from epiphytes in the rainforest near Ko-
lonia, on the Micronesian island of Pon-
ape by M. Sabath on 27 January 1970 (in
the Museum of Comparative Zoology);
allow females of this genus to be de-
scribed for the first time.

Diag^nosis. Females are distin-
guished from those of other uloborid gen-

era by having an epigynum with eight
ventral atria (Fig. 160). The palp of Pu-
rumitra males (Fig. 157), like those of
Zosis (Plate 7-C) and Octonoha (Plate 6-
C) has a tegular spur, but unlike Zosis
they have a broad, concave median
apophysis spur and a small, narrow teg-
ular spur. Their anterior eye row is
strongly recurved such that a line across
the AMEs' posterior margins passes half
an ALE diameter anterior to the ALEs
rather than along the ALEs' anterior bor-
der as in Octonoha. Unlike the males of
either genus, their large median apoph-
ysis bulb is flattened and its lateral sur-
face has a sclerotized ridge running be-
tween two depressions. Purumitra males
are smaller than those of Zosis and Oc-
tonoha, having a carapace length of less
than 0.9 mm.

Description of Type Species. Female
total length 3.0 mm, carapace length 0.8
mm, sternum length 0.6 mm. Male cara-
pace length 0.8 mm, sternum length 0.4
mm. Length and width of female cara-
pace equal, male carapace 0.84 as wide
as long. In lateral view cephalic region of
female carapace level and straight tho-
racic region slopes to petiole. In males
carapace highest just anterior to thoracic
depression, its cephalic region sloping
downward to ocular area and its rounded
thoracic region sloping to petiole. Both
eye rows of females and posterior eye
row of males recurved such that a line
across median eyes' posterior margins
passes through anterior quarter of poste-
rior eyes' diameter (Fig. 159). Anterior
eye row of males more strongly recurved
so that such a line passes half an ALE
diameter anterior to ALE anterior mar-
gin. Median ocular area nearly square, its
length and anterior width four-fifths its
posterior width. Clypeus in anterior view
three-fifths AME diameter. Sternum of
female 0.69 and of male 0.78 as wide as
long. Female palpal femur not flattened.
Female calamistrum half as long as meta-
tarsus IV. Femur I of females 2.0 and of

508

Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

males 1.9 times carapace lenj^th. Abdo-
men teardrop-shaped (Fig. 159) without
tul)ercles, its maximum width and height
half its length and attained midway along
its length. Coloration ol males and fe-
males similar. Carapace very light tan
with broad, brown paraxial stripes ex-
tending its full length (Fig. 159). Tho-
racic region with thin gra\' marginal
stripes. Sternum tan with thin gray mar-
gins. Dorsum of abdomen white with a
gray posterior tip, venter gray with a thin
median longitudinal white stripe, and lat-
eral abdominal surfaces white with a
wide dorsal gray longitudinal stripe and
a broken ventral gray longitudinal stripe.
Numerous fine tracheoles extend into
cephalo thorax.

Male Palpus. No ventral femoral tu-
bercles are visible. Broad cymbium with
two apical macrosetae (Fig. 157). Small
proximal portion of the tarsus includes a
middle hematodocha and gives rise to a
large, somewhat flattened median apoph-
ysis. Retrolateral face of median apoph-
ysis bulb with a heavily sclerotized,
curved central ridge between two depres-
sions (Fig. 157). Short, broad median
apophysis spur with concave retrolateral
depression. Embolus of intermediate
length, supported along its proximal sur-
face by a small tegular spur.

Epigynum. The heavily sclerotized
ventral epigynal surface has a small an-
teromedian and a larger posteromedian
atrium, each divided laterally by a thin
median septiuii (Fig. 160). Also present
are a pair of large, oval, anterolateral atria
and a pair of smaller, triangular, postero-
lateral atria. An opening is located at each
dorsolateral margin of the posterior me-
dian atria and leads via a very short duct
to an oval spermatheca from whose pos-
terior surface a fertilization duct arises
(Fig. 161).

Natural Histonj and Distribution.
Nothing is known of this genus' natiual
history. Its only known species is found
on central Pacific Islands of the Philip-
pine and Caroline groups.

Zosis Walckenaer

Figures 2, 162-174, Plate 7-C; Map 2

Zosis Walckenaer, 1837, Histoire naturelle des In-
sectes, Apteres, Atlas: 12, pi. 20. Type species by
monotypy Zosis caraihe. The genus name is mas-
culine.

Orithi/ia Blackwall, 1858, Ann. Mag. Nat. Hist.,
2(3): 331. Type species by original designation
Orithijia wiUiamsii Blackwall, 1858, ibid.

Dia(i.nosi.s. Zosis males (Fig. 173,
Plate 7-C) are distinguished by a large
tubercle on the proximal, ventral surface
of the palpal femur; a broad, flat tegular
spur; a dome-shaped median apophysis
bulb, and a clawlike median apophysis
spur. Females are distinguished by
broad, flattened epigynal lobes (Figs.
169, 170) which are medially separated,
have posterolateral, triangular sclerites;
and form a pair of posterior atria (Fig.
171) in which openings are found. Cara-
pace (Figs. 162, 163) and leg (Fig. 174)
color patterns are useful in distinguish-
ing members of this genus.

Description. Female carapace width
0.8 carapace length, male carapace width
0.9 length. Female cephalic region rises
only slightly from shallow thoracic
depression, thoracic region flat. Conspic-
uous thoracic groove of males transverse
and top of carapace flat, sloping anterior-
ly with ocidar region lower than thoracic
region. Female cephalic region at ALE
half and male cephalic region 0.4 cara-
pace width (Figs. 162, 163). Female clyp-
eus height in anterior view half and male
clypeus height one AME diameter. An-
terior and posterior eye rows recurved,
posterior such that a line across posterior
margins of PME passes through anterior
half of PLE diameter. Male AMEs on a
mound and anterior eye row more strong-
ly recurved than posterior (Fig. 163). In
females the two rows about equally re-
ciuved (Fig. 162). Median ocular area
nearly square. Sternum width 0.63 times
length (Fig. 164). Prolateral surface of fe-
male palpal femur slightly flattened. Fe-
male first femur 1.8 times and male first
femur 1.5 times as long as carapace. Fe-

Revision of Uloboridae • Opell 509

Figures 157-161. Purumitra grammica (Simon). 157. Prolateral view of male palpus. 158. Dorsal view of female right first
femur. 159. Dorsal view of female. 160. Ventral view of epigynum. 161. Dorsal view of cleared epigynum. Figures 162-
168. Zosis geniculatus (Olivier). 162. Dorsal view of female. 163. Dorsal view of male. 164. Female sternum, endites, and
labium. 165. Dorsal view of female right first femur. 166. Retrolateral view of female left first patella showing lyriform
organs. 167. Prolateral view of male left first femur, patella, and tibia. 168. Dorsal view of female lorum.

male calamistruni half length of metatar-
sus IV. Male first femur with one distal
prolateral and three dorsal macrosetae
(Fig. 167). Tibia I with eight distal pro-
lateral, seven retrolateral, and nine dorsal
macrosetae. Width and height of female
abdomen three-fifths its length, usually
with a single, median tubercle in anterior
third of its dorsum (Fig. 162). Male ab-
domen oval without tubercles, its height

and width half its length (Fig. 163). Col-
oration of members of this genus is dis-
tinctive and given in detail under the
species description. Tracheal system
characterized by small tracheoles extend-
ing to cephalothorax (Fig. 2).

Male Palpus. Proximal ventral sur-
face of femur with one small median and
one large lateral tubercle (Plate 7-C).
Two long setae project from the cym-

510 Bulh'tiu Museum of Comparative Zoology, Vol. 148, No. 10

bium's tip. Middle hematodocha large
and its dome-shaped median apophysis
bulb bears a hooked median apophysis
spur (Fig. 173, Plate 7-C). A broad, 'flat
tegular spur serves as a guide for the em-
bolus, running along its proximal surface
and nearly meeting the median apophy-
sis. The small, flat, unsclerotized lobe
arising from the median apophysis bulb's
center may be homologous with the well-
tormed conductors of PJiiloponeUa, Po-
nella, Mia^rammopes, and Uloborus.

Epigynum. In ventral view two flat-
tened, weakly sclerotized posterior lobes
extend either posteriorly (Fig. 169) or
ventrally (Fig. 170). Posterolateral or
ventrolateral margins of the widely sep-
arated posterior lobes each have a trian-
gular sclerite (Fig. 169). Posterior surface
of eacli lobe forms an atriiun at whose
median margin an epigynal opening is
located (Fig. 171). Dorsally, the two
crypts merge and are bordered by the
straight, ventral posterior plate border. In
posterior view this plate is twice as wide
as high. Duct leading from each epigynal
opening loops twice before entering a
spherical spermatheca from whose pos-
terior surface a fertilization duct arises
(Fig. 172).

Note. The cosmotropical species Zo-
si.s penicillatus is the only member of this
genus I recognize for the New World.
The smaller Uloborus costalimae Mello-
Leitao, 1917 female which has a total
length of 4.0 mm, a carapace length of 1.2
mm, and a median hump on the abdo-
men's dorsum may represent a second
American species. However, the female
holotype (and only specimen discovered
in this study), collected from the Brazil-
ian state of Alagoas by Costa Lima and
housed in the Museu Nacional of Rio de
Janeiro, Brazil, has been dried and is not

in good enough condition to allow a use-
ful redescription of this species.

Natural History and Distribution.
This species appears to have good pow-
ers of dispersal, to be commonly associ-
ated with man-made structures, and to be
quite resistant to desiccation. The latter
two factors suggest that it may have been
introduced into many areas through com-
merce. In the New World this cosmotrop-
ical species of orb-weavers is found from
the Gulf Coast states of the United States
through Central America and the West
Indies, and into the northern two-thirds
of South America.

Zosis geniculatus (Olivier)

Figures 2, 162-174, Plate 7-C; Map 2

Aranea ^eniculata Olivier, 1789, Araignee, Aranea.
in Encycl. nieth., Hist, nat., Inst. Paris, 4; 214.

Zosis caniihe Walckenaer, 1837, Histoire naturelle
des Insectes, Apteres, Atlas: 12, pi. 20. Female
holotype lost.

Uluhorus zosis Walckenaer, 1841, Histoire natu-
relle des Insectes, Apteres, 2: 231.

Uloborus lafrcillei Thorell, 1858, Ofvers. Kongl.
Vet. Akad. Forh., 15: 196.

Orithyia wiUiamsii Blackwall, 1858, Ann. Mag. Nat.
Hist., 2(3): 331. One male, one female, and one
immature syntypes from Amhoina Island, Am-
I)oina (Moluccas) Islands, bottle 161, tube 2, Uni-
versity Museum, Hope Department of Entomol-
ogy, Oxford University, examined.

Uloborus domesticus Doleschall, 1859, Act. Soc.
sci. Ind.-Neerl., 5: 46. Not with other Doleschall
types in Rijksmuseum van Natuurlijke Historie,
Leiden, Holland; presumed lost.

Uloborus williamsii: — O. Pickard-Cambridge, 1871,
Proc. Zool. Soc. London, 1871: 617.

Uloborus Reniculatus: — Thorell, 1890, Ann. Mus.
civ. stor. nat. Genova, 10(2): 291. Bonnet, 1957,
Bibliographia Araneorum, 2: 4762. Muma and
Gertsch, 1964, Amer. Mus. Novitates, 2196: 37.

Zosis geniculatus: — Lehtinen, 1967, Ann. Zool.
Fennici, 4: 277.

Description. Male total length 4.0 to
6.3 mm (X = 4.96, SD = 0.77, N in all
cases = 13). Carapace length 1.6 to 2.5

Figures 169-174. Zosis geniculatus (Olivier). 169, 170. Ventral view of epigynum. 171. Posterior view of epigynum. 172.
Dorsal view of cleared epigynum. 173. Retrolateral view of male palpus. 174. Dorsal view of female left first femur, patella,
and tibia. Figures 175-180. Octonoba species 3. 175. Female carapace. 176. Male carapace. 177. Retrolateral view of
apical region of male left palpus. 178. Ventral view of epigynum. 179. Posterior view of epigynum. 180. Dorsal view of
cleared epigynum.

Revision of Uloboridae • Opell 511

512 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

mm (X = 2.03, SD = 0.23), sternum
length 1.0 to 1.5 mm (X = 1.20, SD =
0.15). Female total length 6.3 to 8.4 mm
(X = 7.21, SD = 0.47), carapace length
2.0 to 2.6 mm (X = 2.35, SD = 0.17), ster-
num length 1.2 to 1.8 mm (X = 1.52, SD =
0.15, N in all cases = 15). Coloration of
males and females similar (Figs. 162),
163). Carapace gray to reddish brown
with light lateral margins, cephalic re-
gion, and posterior median region. Ster-
num tan, often slightly darker in center
than at margins (Fig. 164). Coloration of
all legs similar to first leg (Fig. 174) ex-
cept for four to six dark spots on proximal
region of femur which are found only on
legs I and II. Matatarsi and tarsi brown
with no conspicuous bands. Abdomen
usually white without conspicuous mark-
ings.

Octonoba new genus

Figures 175-185, Plates 6-C, D, 7-D;
Map 1

Type. The type species of this genus is Ulohorus
octonarius Muma, 1945, Proc. Biol. Soc. Wash-
ington, 58: 91. Male holotype and female para-
type from College Park, Maryland, collected 7
July 1943 by \V. F. Jeffers and M. H. Muma, in
the American Musum ot Natural History, exam-
ined. The genus name is an arbitrary combination
of letters and is feminine.

Note. Examined material contained
four distinct Octonoha species, of which
I am able to identify only O. octonaria.
Consideration of the remaining three
Oriental species is necessary for an un-
derstanding of the genus. Designation of
these as species 1-3 is done for clarity
and is not an attempt to name any of these
species. To do so without careful exami-
nation of the types of all Oriental species

likely to belong to this genus would be
unwise. Lehtinen (1967) synonymizes U.
octonarius and U. sinensis Simon, 1880
from China, placing the newly combined
species into the genus Zosis. To this ge-
nus he also adds U. varians Rosenberg
and Strand, 1906 from China and Japan
(which he synonymizes with U. defectus
Rosenberg and Strand and U. duhius
Rosenberg and Strand, 1906), V . sijho-
tides Rosenberg and Strand, 1906 from
Japan, Ar^yrodes yesoensis Saito, 1934
from Japan. Lehtinen also places U. mun-
dior Chamberlin and Ivie, 1936; U. cos-
talima Mello-Leitao, 1917; and U. ursi-
niis Mello-Leitao, 1917 into the genus
Zosis. Lehtinen indicates he did not ex-
amine type specimens of U. octonarius
and I cannot accept his synonymy of U.
octonarius and U. sinensis. Type speci-
mens of Ulohorus ursinus and U. costa-
limae were damaged (the former's abdo-
men lost and the latter dried) and cannot
be treated with certainty. Ulohorus mun-
dior is a synonym of Philoponella repuh-
licana. I agree with Lehtinen that U. oc-
tonarius is more closely related to Zosis
than to any other genus (except possibly
Purumitra) and that it bears a striking re-
semblance to several Oriental species.
However, I believe that these species are
more related to one another than any one
is to Zosis getiicidatus and for this reason
propose Octonoha as a new genus.

Diagnosis. Octonoha males are dis-
tinguished from those of all other ulobo-
rids except Purumitra by having both an
enlarged, concave median apophysis and
a conspicuous hematodocha (Plates 6-C,
D; 7-D). Octonoha males are distin-
guished from those oi Puruniitra by hav-
ing: 1. a carapace length greater than 1.4

Figures 181-182. Octonoba species 3. 181. Retrolateral view of male left first femur, patella, and tibia. 182. Dorsal view
of female right first femur. Figures 183-185. Octonoba species 1. 183. Prolateral view of male left first femur, patella,
and tibia. 184. Ventral view of epigynum. 185 Posterior view of epigynum. Figures 186-191. Daramuliana gibbosa (L.
Kocfi). 186. Female carapace. 187. Lateral view of female carapace. 188. Lateral view of female abdomen. 189. Ventral
view of epigynum. 190. Dorsal view of cleared epigynum. 191. Dorsal view of female right femur. Figures 192-197. Ponella
lactescena (Mello-Leitao). 192. Female carapace. 193. Male carapace. 194. Lateral view of female carapace. 195. Dorsal
view of female. 196. Dorsal view of female right first femur. 197. Lateral view of female abdomen.

Revision of Uloboridae • Opell 513

514 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

mm rather than 0.9 mm or less, 2. first eral, five to eight dorsal, and zero to five
femur no greater than 1.7 (rather than ventral macrosetae. Height and width of
twice) the carapace length, and 3. by hav- female abdomen 0.6 its length, with four
ing a more reduced median apophysis pairs of small paramedian tubercles,
bulb. Octono])(i and Ulohorus females Male abdomen half as wide and two-
ha\ e slender, posterolateral epigynal fifths to three-fifths as high as long. Car-
lobes which are contiguous at their bases apace brown with a thin median light
(Figs. 137, 145, 178, 184). The posterior stripe and a pair of light, marginal, Ion-
eye row of Octonoha (Figs. 175, 176) re- gitudinal stripes. Numerous fine trache-
cuned such that a line across the PMEs' oles extend into cephalothorax.
posterior margins passes along the PLEs' Male Palpus. Femur with two nearly
anterior margins; whereas in Uloboru.s it equal-sized proximal tubercles on a ven-
passes at least one quarter PLE diameter tral mound (Plate 6-A, D). Two stout ter-
anterior to the PLE (Figs. 132, 133). In minal spines extend from cymbium's tip.
OctonoI)a females the median ocular Middle hematodocha large and tegulum
area's anterior width is 0.7 its posterior of two species a spur along embolus'
width; whereas the median ocular area of proximal surface (Fig. 177, Plate 6-C). In
Uloboru.s is square. Octonoha females O. octonaria (Plate 7-D) tegular spur
have a conspicuous atrium (Fig. 179) or small, apparently providing little support
pair of atria on the epigynum's postero- for embolus, but in species 1 where it is
dorsal surface (Fig. 185) which is absent long and grooved (Plate 6-C), tegular
in Ulohorus (Figs. 138, 146). spur probably affords some embolic sup-
Description. Female total length 4.2 port. In O. octonaria and species 1 me-
to 5.2 mm, carapace length 1.5 to 1.6 mm, dian apophysis bulb is a small, convex,
stermnn length 0.9 to 1.0 mm. Male total semicircular sclerite whose spur is en-
length 4.0 to 4.8 mm, carapace length 1.4 larged and has a proximal surface that is
to 1.8 mm, sternum length 0.8 to 1.0. Car- either concave (species 1) (Plate 6-C) or
apace width 0.85 to 0.88 its length. Ce- rolled inward at the edges to form a tube
phalic and thoracic regions flat in lateral (O. octonaria, Plate 7-D). A small irreg-
view, PMEs at the same height as shal- ular, unsclerotized lobe arising from the
low thoracic pit. At ALEs the cephalic median apophysis bulb's center (Plate
regions' width two-thirds maximum car- 7-D) may be homologous with the con-
apace width in females and half in males ductor of Philoponella, Ponella, Mia-
(Figs. 175, 176). Both eye rows recuived grammopes, and Ulohorus. Reduction in
such that a line across median eyes' pos- size of the median apophysis bulb and
terior margins passes along anterior mar- tegular spur appears associated with en-
gin of lateral eyes (Figs. 175, 176). Me- largement of the median apophysis spur,
dian ocular area's anterior width and This is seen when species 1 (Plate 6-C)
length 0.7 its posterior width. In anterior and species 3 (Fig. 177), O. octonaria
view clypeus height of males three-fifths (Plate 7-D) and species 2 (Plate 6-D) are
an AME diameter and of females five- compared. In species 2 (Plate 6-D) where
sixths an AME diameter. Sternum width the median apophysis spur forms a U-
0.65 to 0.72 its length. First femur 1.50 to shaped trough, the tegular spur is lost,
1.65 times carapace length. Female pal- and the median apophysis bulb reduced
pal femur not flattened and calamistrum to a small, flat plate. The embolus of O.
half as long as metatarsus IV. Male first octonaria and species 1 is long and thin,
femur with three to four prolateral, one That of species 2 (Plate 6-D) has a broad
to three retrolateral, and one dorsal mac- base and short central radix,
roseta (Figs. 181, 183). First tibia with Epigynum. The epigynum consists of
11 to 16 prolateral, three to four retrolat- either: 1. a ventral mound with two pos-

Revision of Uloboridae • Opell 515

terolateral lobes, each with a dorsal, cir-
cular atrium at its base (Figs. 184, 185),
or 2. a ventral mound with two posterior
lateral lobes extending over a common
posterior atrium (Figs. 178, 179). The lat-
ter condition is similar to that of Zosis
genicuhitus (Figs. 169-171) and it is in-
teresting to note that the male tegular
spurs oi both species are broad and long.
The convex posterior plate is one-fourth
to three-fourths as high as broad (Figs.
179, 185). A duct leads from each atrium
or from each lateral margin of the com-
mon atrium to a spherical spermatheca
from whose posterior surface a fertiliza-
♦ tion duct arises (Fig. 180).

Natural History and Distribution. Oc-
tonoha octonaria spins an approximately
horizontal orb-web similar to many other
uloborids (Eberhard, in preparation).
This species is known only from the
United States where its range extends
from Maryland through Alabama, north-
central Texas, and eastern Kansas. Species

1 comes from Kunming in the southwest-
ern Chinese province of Yunnan, species

2 from Okinawa, and species 3 from Tai-
wan.

Daramuliana Lehtinen
Figures 186-191; Map 1

Daramuliana Lehtinen, 1967, Ann. Zool. Fennici,
4: 227. Type species by original designation
Ulohorus fiihhosus L. Koch, 1871, Arachn. Aus-
tral., 1: 1, 228. Female holotype from Upolus, Sa-
moa, in Zoologisches Museum, Hamburg, exam-
ined. The generic name is feminine.

Diagnosis. Daramuliana females are
characterized by having: 1. a medially
divided, circular, ventral epigynal atrium
(Fig. 189), 2. a carapace whose width is
0.75 its length (Fig. 186), and an abdo-
men which is 0.38 as wide as long and
has a median, dorsal tubercle in the pos-
terior 0.57 of its length (Fig. 188).

Description of Type Species. Female.
Total length 4.0 to 4.6 mm, carapace
length 1.3 to 1.5 mm, and sternum length
0.8 to 0.9 mm. Carapace width three-
fourths carapace length. Cephalic and

thoracic regions each slope upward from
a shallow thoracic depression (Figs. 186,
187). At ALEs cephalic area half as wide
as carapace. Both eye rows recurved, the
anterior such that a line across AMEs'
posterior margin passes through anterior
quarter of ALEs' diameter and posterior
such that a line across PLEs' posterior
margin passes two-thirds a PLE diameter
anterior to PLEs' anterior margin (Fig.
186). Median ocular area's anterior width
half and its length two-thirds its posterior
width. Clypeus height in anterior view
two-fifths AME diameter. Sternum width
0.42 length. Female palpal femur slightly
flattened. First femur twice as long as car-
apace. Calamistrum half as long as meta-
tarsus IV. Abdomen width and height
0.38 length, median tubercle in posterior
0.57 of the its length (Fig. 188). Fine
tracheoles arising from two stout tracheal
trunks appear restricted to the abdomen.

Epigynum. On the epigynum's ventral
surface is a medially divided circular
atrium (Fig. 189). An epigynal opening is
found at the posteromedian margin of
each atrial half, near the base of a small
scape on the median septvmi's posterior
margin. From each opening a long, thin
duct extends anteriorly and then loops
posteriorly to connect to a spherical sper-
matheca from whose posterior margin a
ferlization duct arises (Fig. 190).

Natural History and Distribution.
Nothing is known of the natural history
of the members of this genus. Its two rec-
ognized species (Lehtinen, 1967) have
been collected on Australia, Samoa, and
on Viti Levu in the Fiji Islands.

Ponella new genus

Figures 192-202; Map 1

Tijpe. Type species Uloborus lactescens Mello-
Leitao, 1947, Boletim do Museu Nacional (N.S.),
Rio de Janeiro, Zoologia, No. 80: 4. The genus
name is an arbitrary combination of letters and is
feminine.

Diagnosis. Members of this genus
have a total length of less than 3.0 mm.
Ponella (Fig. 193), Zosis (Fig. 163), and

516 Bulletin Museum of Cumparative Zuology, Vul. 148, Nu. 10

Philoponella (Figs. 205, 217) males have slightly anterior to its center (Figs. 195,

an oval carapace with a wide, transverse 197). Male abdomen oval without humps,

thoracic groove and AMEs on a small its width and height two-thirds its length,

mound. Ponella males are distinguished Fine tracheoles extend into the cephalo-

l)y ha\'ing a long, narrow conductor thorax.

which lies along the embolus' distal sur- Male Palpus. Femur with two proxi-

face, lacks a basal lobe, and has a slender, mal, ventral processes; a large lateral one

distal branch (Fig. 201). First tibia of and a smaller median one. No conspicu-

males (Fig. 202) has three prolateral, ous macrosetae visible on distal cymbial

three retrolateral, and four dorsal macro- rim. Middle hematodocha well devel-

setae — fewer than either Zosis or Philo- oped giving rise to a median apophysis

ponella. Ponella females are distin- consisting of a flat bulb with a large lat-

guished by having a pair of low, widely eral hooked spur (Fig. 201). Conductor

separated, ventral epigynal mounds an- with a short cylindrical proximal lobe and

terior to which a pair of openings is found a long, thin distal branch about five times

(Figs. 198, 199). as long as wide and with proximal surface

Description. Carapace about 0.9 times trough-shaped to receive the embolus,
as wide as long. Male thoracic groove Epigynum. In ventral view (Fig. 198)
deep and transverse and carapace dorsal- two light, slightly raised posterior lobes
ly flat (Fig. 193), slanting forward so that appear near epigynum's posterior edge,
cephalic region is much lower than tho- Anterior to these is a pair of small, incon-
racic region. Transverse female thoracic spicuous openings. In posterior view
groove narrow but conspicuous (Figs. (Fig. 199) a posterior plate extends be-
192, 195), cephalic and thoracic regions tween the lobes and bears a pair of small
about equal in height (Fig. 194). At level midlateral bumps. From each opening a
of ALEs female cephalic region 0.70 duct makes four loops before connecting
times and male cephalic region 0.60 with a large, thick-walled spermatheca
times carapace width. Male AMEs on a from whose posterior margin a fertiliza-
small, prominent, anteriorly directed tion duct arises (Fig. 200).
mound (Fig. 193), but in females this Natural History and Distribution. The
mound is not distinct (Fig. 192). Both eye one described species of this genus has
rows recurved, the posterior only slightly been collected from Paraguay and south-
so such that a line across posterior mar- ern Brazil. One female was collected
gins of PMEs passes through center of with a chain of three flat, stellate eggsacs,
PLEs. Such a line across AMEs' posterior each about 4.0 x 2.8 mm.
margins passes across anterior margins of

ALEs of females and in males 1 to 1.5 Ponella lactescena (Mello-Leitao),

ALE diameters anterior to ALEs. In an- new combination

terior view female clypeus height half an Figures 192-202; Map 1
AME diameter and in males two AME

diameters. Median ocular area of females ^'/'^f^'""^ '"''""■';' ^c'? I' ";^'''?"; ^^'^'^' ^^'''*!'" '^°
1 ,• , 1- I .1 1 .1 Museu Nacional (N.b.), Kio de laneiro, Zoologia,
sciuare and of males slightly longer than n„ gO: 4. Immature female and cne mature male
Wide. Sternum width 0.67 its length. syntypes from Carmu do Rio Claro in the Brazil-
First femur 1.1 times as long as carapace ian state of Minas Gerais, collected by J. C. M.
in females and in males equal to carapace Cai-valho, in the Museu Nacional of Rio de Ja-
length. Female calamistrum half as long "^"""' ^'^'"*'' ^^^^"""^^^d-

as metatarsus IV. Male first femur with Description. Male. Total length 2.8

one dorsal, two distal prolateral, and two mm, carapace length 1.3 mm, sternum

distal retrolateral macrosetae. Female ab- length 0.7 mm. Fourth coxae separated

domen half as wide and two-thirds as by a distance equal to the width of each,

high as long, with two dorsal humps at or Carapace tan with no conspicuous color

Revision of Uloboridae • Opell 517

204

republicana

204

P. divisa

P. tingena

203

206

Figures 198-202. Ponella lactescena (Mello-Leitao). 198. Ventral view of epigynum. 199. Posterior view of epigynum.
200. Dorsal view of cleared epigynum. 201. Retrolateral view of male left palpus. 202. Prolateral view of male left first
femur, patella, and tibia. Figure 203. Apical views of left median apophysis spurs of Phlloponella republicana species
group. Figures 204-206. Phlloponella republicana (Simon). 204. Dorsal view of female. 205. Dorsal view of male. 206.
Dorsal view of female left femora, showing position of trichobothrial rows.

518 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

markings. Steniuni tan with li.uht gray an-
terior margin and center. Dorsum of ab-
domen light with flecks of" white and with
light gray posterior tip. Venter with gray
patch between booklung covers and a
broad, light gray median area. Legs tan.

Female. Total length 3.0 to 3.5 mm,
carapace length 0.9 to 1.1 mm, sternum
length 0.7 mm. Carapace light gray with
narrow white border, lighter ocular re-
gion, and darker patch at posterior ce-
phalic region (Fig. 195). Eyes encircled
by dark gray rings. Sternum dark gray.
Dorsum of abdomen light with no mark-
ings except gray posterior tip in some
specimens. Venter dark gray with two
think widely spaced white paraxial stripes
running its entire length. Legs with no
conspicuous color markings.

Distribution. Eastern Paraguay and
southern Brazil.

Philoponella Mell-Leitao

Figures 1, 203-288, Plates 6-E, 7-B;
Maps 3-5

Philoponella Mello-Leitao, 1917, Arch. Esc. Sup.
A^r. Med. V^eter., 1(1): 8. Type species by original
designation and monotypy Ulohorus republica-
nus Simon, 1891, Ann. Soc. ent. France, 60: 8. The
genus name is feminine.

Note. Until Lehtinen, 1967, Ann.
Zool. Fennici, 4: 258, resurrected this ge-
nus, the name Philoponella was not rec-
ognized by anyone other than its author.
However, its first formal synonymy with
Ulohorus was by Roewer, 1954, Katalog
der Araneae, 2(b): 1344. This genus cor-
responds with the Ulohorus republica-
nus species group Muma and Gertsch,
1964, Amer. Mus. Novitates, 2196: 29. In
addition to the type species, U. repuJAi-
canus, this species group contained U.
variegatus, U. oweni, U. arizonicus, and

"a series of similar species from the
Americas."

Diagnosis. PJiiloponella males are
distinguished by having: 1. an oval cara-
pace with a broad, transverse thoracic
groove (Figs. 205, 217), 2. a nearly
straight posterior eye row with about
equally spaced eyes, 3. a clypeus height
0.6 to 1.5 an AME diameter, 4. a well-
developed conductor spur (Plate 7-B),
and 5. a first tarsus with four to five pro-
lateral, four to five retrolateral, and six to
eight dorsal macrosetae. Females are
characterized by having: 1. a nearly
straight posterior eye row with about
equally spaced eyes (Figs. 204, 216), and
2. a ventral epigynal atrium whose pos-
terior margin is formed by two lateral
lobes (Figs. 208, 239, 250, 287).

Description. Carapace width 0.9 to
1.0 mm long. At ALEs cephalic region of
females gradually narrowed to about half
maximum carapace width; that of all
males except those of P. tingena less
conspicuously narrowed, making their
carapace oval (Figs. 205, 217). Cephalic
region rises slightly from a narrow trans-
verse or semicircular thoracic groove to
the ocular area. The AMEs on a low an-
teriorly directed mound, more prominent
in males than in females. Eyes in two re-
curved rows, posterior row only slightly
curved so that a line across PMEs' pos-
terior margins passes through or posterior
to the PLEs' centers (Fig. 204). Median
ocular area is either square or 0.8 as wide
anteriorly as posteriorly. The AME-ALE
separation two to three times AME sep-
aration and PME-PLE separation 0.6 to
1.0 times PME separation. Clypeus
height in anterior view 0.7 to 1.5 times
an AME diameter. Sternum width 0.67
length. Endites 1.3 and labium 1.1 times

Figures 207-215. Philoponella republicana (Simon). 207. Lateral view of epigynum. 208. Ventral view of epigynum. 209.
Posterior view of epigynum. 210. Dorsal view of cleared epigynum. 211. Dorsal view of female first femur. 212. Retrolateral
view of left embolus (black), sperm reservoir, and fundus. 213. Ventral view of male, left palpal femur. 214. Retrolateral
view of male left palpus. 215. Retrolateral view of expanded male left palpus. Figures 216-219. Philoponella divisa n.
sp. 216. Dorsal view of female. 217. Dorsal view of male. 218. Retrolateral view of male left palpus. 219. Apical view of
male left palpus.

Revision of Uloboridae • Opell 519

520 Bulletin Museum uf Comparative Zoolugij, Vol. 148, No. 10

as long as wide. Female palpal femur not 250, 275, 287) or eomprised of two dis-
flattened, ealamistrum hali as long as tinet lobes joined medially (Figs. 208,
fourth metatarsus. Male first femur with 239). Openings may he at the atrium's
three prolateral, one or two retrolateral, eenter (Fig. 250) or in the atrium's pos-
and one or two dorsal maerosetae. Male terior half (Figs. 208, 230, 266) and are
first tibia with four or five prolateral, four separated by a distanee of lialf (Fig. 208)
or five retrolateral, and six or eight dorsal to 0.8 (Fig. 250) the epigynum's width,
maerosetae. Abdomen width and height Anterior and posterior rims may be con-
one-third to one-half its length. In fe- tiguous (Figs. 250, 259, 275) or separated
males it attains its maximum width about laterally by wide troughs (Figs. 208, 221,
midwa>' along its length and in males in 239). Epigynal duets make one or two
the anterior third of its length. Abdomen loose 180° loops (Figs. 223, 268) or spiral
height usiialK' attained at or slightly an- tightly around themselves (Figs. 252,
terior to widest area. Fine traeheoles 277) before connecting to a pair of sper-
which arise from two stout tracheal mathecae from whose posterior margins
trunks are restricted to the abdomen fertilization ducts lead. The republicana
(Fig. 1). species group appears more primitive

Male Palpus. Femur's ventral surface than the seniiphimosa group. In semiplu-

with one or two small, proximal tubercles mosa females there is more extensive

(Figs. 213, 269). Tarsus with two long, coiling and spiraling of ducts (Fig. 252),

apical cymbial setae and a large middle more complete median fusion and an-

hematodocha (Plate 7-B). A grooved con- terolateral displacement of epigynal

ductor blade arises near the proximal, openings and closure of lateral epigynal

ventral border of the median apophysis troughs. Members of the /fl.sc/afr/ species

bulb and continues as a narrow, heavily group have coiled epigynal ducts (Fig.

sclerotized conductor spike into the prox- 277) and closed epigynal troughs (Figs.

imity of the median apophysis spur. Prox- 275, 287) indicating they are intermedi-

imal conductor lobe may be nearly ate between the two other species

s(iuare, weakly sclerotized, and often dif- groups.

ficult to see as in the republicana group Natural History and Distribution.

(Figs. 214, 215; Plate 7-B) or it may be Members of many species in this genus

longer than broad, heavily sclerotized, are found in colonies consisting of nu-

and conspicuous as in the vittata (Fig. merous orb-webs built in a common, ir-

270) and semiplumosa (Fig. 253) groups, regular framework. Some webs are asso-

Epigynvim. Females are character- ciated with those of spiders belonging to

ized by a well sclerotized epigynum with other families (see family Natural History

a large, deep, central atrium (Figs. 208, section). In the New World, members of

239, 250, 287) bordered by precipitous this genus are found from the extreme

anterior (except in P. vittata. Fig. 266) southern central United States southward

and posterior rims. Posterior rim bears an as well as in the West Indies (Maps 3-5).

epigynal opening at each anterior lateral The genus is also represented in Oriental

margin and may be continuous (Figs, and Australian regions.

Figures 220-223. Philoponella divisa n. sp. 220. Lateral view of epigynum. 221 . Ventral view of epigynum, 222. Posterior
view of epigynum. 223. Dorsal view of cleared epigynum. Figures 224-232. Philoponella tingena (Cfiamberlin and Ivie).
224. Dorsal view of male. 225-227. Dorsal view of female. 228. Lateral view of female carapace. 229. Lateral view of
epigynum. 230 Ventral view of epigynum. 231. Posterior view of epigynum. 232. Retrolateral view of left male palpus.
233. Eggsac. Figure 234. Philoponella signatella (Roewer), dorsal view of female.

Revision of Uloboridae • Opell 521

522

Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Key to Puiloposelu\ Females

1. Clypeus height 0.5 to 0.7 AME diameter;
anterior rim of epigynal atrium liroad,
roimded, and not precipitous (Fig. 266);
epigynal openings in posterior sixth of
atrium; South America (Map 5) vittata

- Clypeus height e(iual to AME diameter;
anterior rim of atrium precipitous and
usually narrow (Figs. 208, 239, 250, 275);
epigynal openings in anterior half (Figs.
242, 250, 287) to posterior third (Figs.
208, 239) of atrium or, if more posterior,
posterior rim projects conspicuously ven-
trally and has a length ecjual to that of the
anterior rim (Figs. 284, 286) 2

2(1). Epigvnal openings separated In- half or
less the atrium's width (Figs. 208, 230,
242); precipitous anterior and posterior
rims separated by conspicuous lateral or
posterolateral troughs (Figs. 208, 221,
239); posterior rim of epigynum 0.4 to 0.6
the epigynum's width and composed of
two lateral lobes separated by a median
depression or notch (Figs. 209, 222, 231,
240); {repuJ)Ucana species group) 3

- Epigynal openings separated by at least
three-fourths atrial width (Figs. 250, 259,
275, 287), or if less, posterior epigynal
plate as high as wide (Figs. 284, 286); an-
terior and posterior rims not separated by
wide, lateral troughs (Figs. 250, 259,
275); posterior rim at least three-fourths
as wide as epigynal atriiun and without
a deep median indentation or notch
(Figs. 250, 251, 275, 276, 287, 288) 7

3(2). Epigynal atriiun with a thin, median, lon-
gitudinal septum (Fig. 221); posterior
margin with a shallow notch separating
lobes (Fig. 222); epigyninn's posterior
face with a middle ridge (Figs. 220, 222);
abdominal dorsum with two broad, light

paraxial stripes (Fig. 216); Colombia

divisa

- Epigynal atrium without a septum (Figs.
208, 239); posterior margin with a deep
notch (Figs. 209, 240) or a broad depres-
sion separating lobes (Fig. 23L); epigy-
num's posterior face without a ventral rim
(Figs. 207, 209, 229, 231, 238, 240); ab-
dominal coloration otherwise (Figs. 204,
226, 234) 4

4(3). Epigynal atrium shallow with broad, lat-
erally directed troughs (Figs. 208, 242);
anterior rim and posterior lobes separat-
ed by a distance ecjual to half the atrial
width (Figs. 208, 242); epigynal openings
visible in ventral view (Fig. 208, 242) ._ 5

- Epigynal atrium deep with posteriorly-
directed troughs (Figs. 230, 239); anterior
rim and posterior lobes sc>parated by a
distance e(jual to 0.30 the atrial width
(Figs. 230, 239); epigynal openings locat-

ed at the anterior margins of the posterior
lobes and hidden in ventral view (Figs.
230, 239) 6

5(4). Epigynal openings on raised mounds in
posterior third of atrium (Figs. 207, 208);
total body length more than 5.4 mm; ster-
num usually dark gray and more than 1.1
mm long; abdominal dorsum with two
pairs of light, lateral, longitudinal, di-
verging white stripes (Fig. 204); Panama
and South America (Map 3) __ republicana

- Epigynal openings in depressions at
atrium's center (Fig. 242); total body
length less than 4.2 mm; sternum tan
with gray posterolateral margins and less
than 0.8 mm long; abdominal dorsum
with a broad, light median stripe (Fig.
241); southern Mexico vicina

6(4). Posterior epigynal lobes widely separat-
ed by a shallow depression (Fig. 231);
anterior epigynal rim one-fifth as long as
atrium (Fig. 230); total body length less
than 4.0 mm; sternum less than 0.8 mm
long; Costa Rica, Panama, and Colombia
(Map 3). tingena

- Posterior epigynal lobes closely spaced
and separated by a deep notch (Fig. 240);
anterior epigynal rim half to a third the
atrial length (Fig. 239); total body length
more than 4.0 mm; sternum length more
than 0.9 nun; Honduras through eastern
Mexico (Map 3) signatella

7(2). Epigynal atrium slightly longer than
wide (Fig. 287); epigynal openings in an-
terior half of atrium (Fig. 287); posterior
face ol epigymun with two lateral grooves
(Fig. 288); northeastern Colombia ._ belhi

- Epigynal atrium about twice as wide as
long (Figs. 250, 259, 275, 284); epigynal
openings at center of atrium (Figs. 250,
275) or in posterior half (Figs. 259, 284);
posterior face of epigvnum without lat-
eral grooves (Figs. 251, 262, 276, 286) 8

8(7). Posterior face of epigynum as high as
broad (Fig. 286); epigynal openings in
the posterior quarter of atrium (Fig. 284);
venter of abdomen usually with a dark W
mark on a light field (Fig. 282); Para-
guay para

- Posterior face of epigynum about twice
as wide as high; epigynal openings at or
slightly behind the atrium's center; ven-
ter of abdomen gray with light paraxial
stripes 9

9(8). Anterior atrial rim continues to the epig-
ynum's posterior margin and encloses the
broad, dorsally sloping lateral regions of
the posterior rim (Fig. 275); large, longi-
tudinal epigynal openings separated by
a distance etiual to 0.6 the atrial width;
Paraguay and southeastern Brazil (Map
5) fasciata

Revision of Uloboridae • Opell 523

- Anterior and posterior atrial rims end
abruptly upon meeting at or slightly pos-
terior to the atrium's center (Figs. 250,
257, 259); small, transverse epigynal
openings are separated by a distance
equal to 0.8 the atrial width; southwest-
ern United States through Central Amer-
ica and the West Indies (Map 4); (semi-
phimosa species group) 10

10(9). Abdominal venter gray with two narrow,
paraxial white stripes (Fig. 249) setose
field anterior to cribellum smaller than
cribellum; lengths of anterior and poste-
rior epigynal rims equal (Fig. 250); ducts
of most specimens do not show through
atrial integument; total length less than
4.5 mm, sternum length less than 1.0 mm,
southeastern Texas through Central
America and the West Indies (Map 4)
semiplumosa

- Abdominal venter with two broad white
stripes (Fig. 256); setose field equal to or
larger than cribellum; anterior epigynal
rim one quarter the length of the poste-
rior rim (Figs. 257, 259); ducts of most
specimens visible through atrial integu-
ment; total length greater than 4.6 mm,
sternum length more than 1.0 mm 11

11(10). Epigynal atrium less than 0.5 mm wide
and 0.4 as long as wide (Fig. 257); epig-
ynum in posterior view without ventro-
lateral gray lines; most specimens darkly
colored (Fig. 255); carapace less than 2.0
mm long; sternum less than 1.4 mm long;
southwestern United States and northern
Mexico (Map 4) oweni

- Epigynal atrium more than 0.6 mm wide
and 0.6 as long as wide (Fig. 259); pos-
terior face of epigynum with dark lines
(Fig. 262); most specimens with no dark
color markings (Fig. 260); carapace more
than 2.3 mm long, sternum more than 1.5
mm long; Arizona (Map 4) arizonica

Key to Philoponella Males
(Males of P. vicina and P. bella are unknown.)

Carapace high and cylindrical (Figs. 278,
279), transverse thoracic groove deep
(Fig. 278), clypeus height twice AME di-
ameter; palpal femur with a retrolateral
tubercle whose length is three times its
basal width (Fig. 281); median apophysis
very small and flat (Fig. 208); Paraguay

and southern Brazil (Map 5) fasciata

Carapace low with rounded margins; tho-
racic groove usually not deep (Figs. 205,
217); clypeus height equal to AME di-
ameter; palpal femur's retrolateral tuber-
cle, if present with a length equal to or
less than its basal width (Fig. 269); me-

dian apophysis bulb not so reduced (Figs.

214, 235, 2.53, 270) 2

2(1). Conductor's proximal lobe at least three

times as long as broad and easily seen
although thin (Figs. 2.53, 270), at least 1.5
times as long as either the conductor
spike or median apophysis spur's apical
projection; median apophysis spur's api-
cal (darkest) projection more robust, its
length equal to or less than twice its basal
width (Fig. 245); median apophysis
spur's basal region surrounded jjy bulb
along half of its perimeter (Fig. 245) 3

- Conductor's proximal lobe small, square,
and usually difficult to see (Figs. 214,

215, 218, 232, 236), no longer than the
conductor spike or median apophysis
spur's apical projection (or conductor
lobe short and with a long, thin, apical
projection iimning parallel to the con-
ductor spur. Fig. 283); median apophysis
spur's apical (darkest) projection long and
slender, about three times as long as its
basal width (Figs. 203, 272); median
apophysis spur's basal region surrounded
by bulb along one quarter or less of its
perimeter (Figs. 203, 272) 6

3(2). Conductor lobe large (Fig. 270); median
apophysis bulb dome-shaped (Figs. 270,
271); dorsum of abdomen with three light
chevrons (Fig. 265), venter with a light
chevron midway between spinnerets and
epigastric furrow; South America (Map
5) vittata

- Conductor lobe small (Fig. 253); bulb
flattened (Fig. 2.53); abdomen not so
marked (Figs. 248, 258, 261); southwest-
ern United States through Central Amer-
ica and West Indies 4

4(3). First tibia with eight or fewer dorsal mac-
rosetae; carapace less than 1.4 mm long,
sternum less than 0.8 mm long; south-
eastern Texas through Central America
and West Indies (Map 4) semiplumosa

- First tibia with nine to 12 dorsal macro-
setae; carapace more than 1.6 mm long,
sternum more than 0.9 mm long; south-
western United States and northwestern
Mexico 5

5(4). Sternum dark; usually nine or ten dorsal
spines on first tibia; carapace less than 1.8
mm long, sternum less than 1.1 mm long;
palpal median apophysis bulb less than
0.6 mm wide; median apophysis spur's
apical (darkest) projection 1.5 times as
long as its basal width (Fig. 245); Map
4 oweni

- Sternum light; usually 12 dorsal spines
on first tibia; carapace more than 2.0 mm
long, sternum more than 1.1 mm long;
palpal median apophysis bulb more than
0.6 mm wide; median apophysis spur's

524 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

apical (darkest) projection two times as
long as its basal width (Fig. 245); Map

4 arizonica

6(2). Conductor lobe with a long, slender api-
cal projection which extends parallel to
and is al)out as long as the conductor
spike (Fig. 283); median apophysis bulb
with a small, apical lobe (Fig. 283); me-
dian apophysis spur's basal region with
a long distal lobe (Fig. 272); first tibia
with four dorsal macrosetae; Paraguay _.
para

- Conductor lobe square, without an apical
projection (Plate 7-B; Figs. 214, 215,
218); median apophysis bulb without an
apical lobe (Figs. 214, 218, 232, 235);
spur's basal region without such a lobe
(Fig. 203); first tibia with si.\ to eight dor-
sal macrosetae {repuhlicana species
group, in part) 7

7(6). In apical view median apophysis bulb
with a lobe which extends along the me-
dian surface of the median apophysis
spur's basal region (Fig. 203); sternum
usually dark gray 8

- Median apophysis bulb without such a
lobe (Fig. 203); sternum tan to reddish
brown with gray posterolateral margins 9

8(7). Total length more than 3.6 mm, sternum
length more than 0.9 mm; median apoph-
ysis bulb diameter 0.4 mm; Panama and
South America (Map 3) repuhlicana

- Total length less than 3.4 mm, sternum
length less than 0.9 mm; median apoph-
ysis bulb diameter 0.3 mm; Colombia
(Map 3) divisa

9(7). Carapace more than 1.2 mm long, ster-
num more than 0.7 mm long; median
apophysis bull> rounded-rectangular (Fig.
235), diameter 0.4 to 0.5 mm; carapace
light tan with two anteriorly diverging
longitudinal gray stripes (Fig. 237); east-
ern Mexico through Honduras (Map 3)
signatella

- Carapace less than 1.2 mm long, sternum
less than 0.7 mm long; median apophysis
bulb semicircular (Fig. 232), diameter 0.2
to 0.3 mm; carapace gray with either no
color markings or with a wide median
light longitudinal stripe (Fig. 224); Costa
Rica through Colombia (Map 3)

tingena

The Philoponella repuhlicana
Species Group

This ^roup contaiii.s P. re))uhlicana, P.
divisa, P. tingena, P. signatella, and P.
vicina. It does not correspond to the
Ulohorus repuhlicana species group of
Muma and Gertsch (1964) which is a syn-
onym of the genus Pliiloponella.

Description. Clypeus height of males
and females equal to AME diameter.
Epigynal openings in posterior third
(Fig. 230) to half (Fig. 208) of epigynal
atrium and seaparated by a distance
equal to half the atrium's width. Anterior
and posterior atrial rims widely separat-
ed, forming broad lateral or posterolateral
troughs (Figs. 208, 221, 239). Anterior
rims precipitous and posterior rim com-
posed of two lateral lobes separated by a
depression or notch (Figs. 208, 209, 239,
240). Each female connecting duct makes
a single loop before connecting with a
spermatheca (Fig. 210). In P. vicina each
duct connects to a small bulb v.^hich, in
turn, connects with a spermatheca (Fig.
244). Conductor blade of male palpus
short and square, about as high as long
(Plate 7-B; Figs. 215, 218). Conductor
spike shorter than slender median apoph-
ysis spur's apical projection (Figs. 232,
235, 236). Prolate ral palpal femoral tu-
bercle slightly larger than retrolateral tu-
bercle and with a length slightly less than
its basal width (Fig. 213).

Philoponella repuhlicana (Simon),
new combination

Plate 7-B; Table 4; Figures 1, 203-215;

Map 3

ulohorus repuhlicanus Simon, 1891, Ann. Soc. ent.
France, 60: 8. Five male and 56 female syntypes
from the forests of San Esteban (6 km S of Puerto
Cabello), Venezuela, collected by E. Simon on 29
February to 27 March 1888, in Museum National
d'Histoire Naturelle, Paris, examined. Bonnet,
1957, Bibliographia Araneorum, 2: 4768.

Ulohorus cuminamensis Mello-Leitao, 1930, Sep.
Arch. Mus. Nac, Rio de Janeiro, 32: 61. Female
holotype (No. 1216) from Cumina [Cuminapane-
ma] River in the Brazilian state of Para, in Museu
Nacional, Rio de Janeiro, Brazil, examined, NEW
SYNONYMY.

Ulohorus mundior Chamberlin and Ivie, 1936, Bull.
Univ. Utah, 27(5): 11. Female holotype, three
male and ten female paratypes from Barro Colo-
rado Island, Panama, collected by A. M. Chick-
ering in 1928, in the American Museum of Nat-
ural History, examined, NEW SYNONYMY.

Zosis mundior: — Lehtinen, 1967, Ann. Zool. Fen-
nici, 4: 277.

Note. Included with U. repuhlicanus

Revision of Uloboridae • Opell 525

• P signatella
▲ P tingena
o P republicana

A Anchicaya River Valley
B El Queremal
C, Call

Valle del Cauca Dept., Colombia

50 km

Map 3. Distribution of Philoponella signatella, P. tingena, and P. republicana (left) and Colombian P. tingena (right).

syntypes were four male and two female
Philoponella servula (=F. vittata). Si-
mon clearly separated these two species
in 1892 [1893]. As he collected both
species at San Esteban during the same
time period it is likely that these P. ser-
vula represent specimens he overlooked
when sorting his collection. A female
Leucauge and male Nephila were also
included with the syntypes.

Diagnosis. Philoponella republicana
females are distinguished by having a
long, shallow, undivided epigynal atrium
whose openings are situated on raised
mounds and easily visible in ventral view
(Fig. 208). Posterior lobes are low and
wider than long (Figs. 208, 209). Females
have a characteristic abdominal color pat-
tern featuring two pairs of diverging
white stripes (Fig. 204). Males are distin-
guished from all others of this species
group but P. divisa by the presence of a
median apophysis lobe which, in apical
view, extends along the median surface
of the median apophysis spur's basal re-
gion (Fig. 203). PJiiloponella republicana
is larger than P. divisa, having a total

length of more than 3.6 mm and a ster-
num length of more than 0.9 mm.

Description. Female. Total length 5.5
to 7.6 mm (X - 6.35, SD = 0.47,_N = 40),
carapace length 1.7 to 2.3 mm (X = 1.96,
SD = 0.12), sternum length 1.1 to 1.4 mm
(X = 1.26, SD = 0.08). Abdomen half as
wide as long with a single broad, raised
area in anterior fourth (Fig. 204). Dark
forms with only faint light carapace and
abdominal markings (Fig. 204). Lighter
forms with light, trilobed carapace mark-
ing whose central lobe extends to front of
carapace. Sternum black. Dorsum of ab-
domen with two or three recurved, nearly
paraxial white chevrons; tip dark gray.
Venter of abdomen with dark gray median
area. Legs reddish brown or light gray
with central light ring on femora and
tibiae. Epigynal atrium open laterally,
about half as long as wide (Fig. 208).
Each opening on a raised area and easily
visible in ventral view (Figs. 207, 209).
Anterior epigynal rim about one-tenth as
long as atrium. Ducts at their closest
point separated by two diameters (Fig.
210).

526 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

Male. Total length 3.6 to 4.6 mm (X =
4.10, SD = 0.18, N = 40), carapace length
1.6 to 1.9 mm (X = 1.72, SD = 0.08), ster-
num length 0.9 to 1.1 mm (X = 0.98, SD =
0.04). Carapace light reddish brown dust-
ed with gray (Fig. 205). Median light area
in the shape of a posteriorly directed ar-
rowhead extending full length of cara-
pace. Sternum dark gray. Dorsum of ab-
domen tan, white pigment spots forming
three fairly distinct che\'rons, tip dark
gray (Fig. 205). Venter of abdomen with
median dark gray area about half as wide
as abdomen. Legs light reddish brown.
The palpal median apophysis bulb has a
width of about 0.4 mm (X = 0.38, SD =
0.02, N = 40) and is crescent to semicir-
cular shaped (Figs. 214, 215).

Natural History and Distribution.
Foimd in Panama and South America
(Map 3). The presocial nature of this
species has been known since Simon's
(1891) description and illustration of P.
republicana from Venezuela (see family
Natural History).

Philoponella divisa n. sp.

Plate 8-A-C; Table 5; Figures 216-223;
Map 3

Types. Female holotype, one male and two female
paratypes in the Museum of Comparative Zool-
ogy, one male and one female paratype in the
American Museum of Natural History. All speci-
mens from the Anchicaya River Valley near the
Central de Anchicaya Limitada, Planta Hidrauli-
ca in the ColomI)ian department of Valle del Cau-
ca, collected 7 to 9 August by B. Opell. The spe-
cific epithet refers to the lateral division of the
epigynal atriuTu.

Dia<i,nosis. Females are distin-
guished by their divided epigynal atrium
(Fig. 221) and dorsal abdominal color-
ation (Fig. 216). Males are distinguished
from all others of this species group but
P. republicana by the presence of a large
median apophysis lobe which, in apical
view, extends along the median surface
of the median apophysis spur's basal re-
gion (Fig. 203). Philoponella divisa
males are smaller than those of P. repub-
licana, having a total l(Migth of 3.4 mm or

less and a sterniun length of 0.9 mm or
less.

Description. Female. Total length 3.0
to 3.6 mm (X = 3.35, SD = 0.23, N = 9),
carapace length 1.0 to 1.1 mm (X = 1.04,
SD = 0.04), sternum length 0.7 to 0.8 mm
(X = 0.72, SD = 0.04). Abdomen width
and height about 0.6 its length with two
dorsal humps in the anterior third of its
length. Most specimens have a dark gray
carapace with light posterolateral mar-
gins and a few have a light posterome-
dian longitudinal stripe (Fig. 216). Ster-
num dark gray to black. Abdomen dark
gray to black with a pair of broad, light
paraxial stripes on dorsum (Fig. 216) and
venter. First femur dark gray with a nar-
row white ring in distal half of its length.
Second through fourth femora gray with
a white ring in proximal third and distal
two-thirds of its length. Patellae dark
gray. Tibiae gray with proximal and cen-
tral white rings. Metatarsi and tarsi gray
with narrow proximal white rings. In ven-
tral view epigynal crypt about 0.2 as
long as wide, divided by a thin median
septum, and with low, closely spaced
posteromedian lobes (Figs. 221). In pos-
terior view raised posterior epigynal face
with two lateral posterior lobes separated
by a shallow notch (Fig. 222). A simple
straight duct leads from each inconspic-
uous posteromedian epigynal opening to
a spherical spermatheca (Fig. 223).

Male. Total length 2.6 to 3.4 mm (X =
3.02, SD = 0.22, N = 8), carapace length
1.3 to 1.4 mm (X = 1.38, SD = 0.35), ster-
num length 0.7 to 0.8 mm (X = 0.78, SD =
0.04). Carapace gray with a light median
stripe and light posterolateral margins
(Fig. 217). Abdomen tan with one or two
mid dorsal paraxial spots and a gray pos-
terior tip (Fig. 217). Legs white to tan,
markings when present, of low contrast.
First and second femora light gray with
a narrow white ring in distal half of its
length. Second through third femora tan.
First, third, and fourth patellae tan, sec-
ond patella light gray. First, second and
fourth tibiae light gray with a narrow,
central, white ring. Third tibia tan with

Revision of Uloboridae • Opell 527

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528 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

a distal gray band. Metatarsi and tarsi
tan. Median apophysis bulb semicircular
(Fig. 218), length and width about 0.3
mm (X = 0.29 mm, SD = 0.01, N = 8).

Ndtural History and Distribution.
Known only from the type locality in Co-
lombia (Map 3). Specimens were collect-
ed from vegetation and rocks of roadside
banks where their webs were always
found adjacent to and connected with di-
plurid funnel webs. Several P. divisa
webs were usually found together, but
colonies did not exceed four individuals.
Webs of immatiues and mature females
were diagonal to vertical (Plate 8-A-C;
Table 5) and featured a signal line run-
ning from the hub to a peripheral sup-
port. The spider hung from this line near
the support and moved to the hub only
when prey vibrations were detected. As
with other P}\iloponeUa species, males
were observed hanging from framework
threads of female and immature orb-webs.

Philoponetla tingena
(Chamberlin and Ivie),
new combination

Plate 8-E-F; Tables 4, 5; Figures 224-

232; Map 3

Uloborus tingens Chamberlin and Ivie, 1936, Bull.
Univ. Utah, 27(5): 10. Female holotype and six
female paratypes hom Bano Colorado Island,
Panama, Canal Zone, eollected by A. M. Chick-
eriny in 1928, in the American Museum of" Nat-
ural History, examined.

Di(i<i,n()si.s. PJiiloponella tin^cna and
P. si<i.n(itell(i are similar. Females of the
former have a broad, shallow depression
separating posterior epigynal lobes (Figs.
230, 231); whereas the latter has a deep
notch separating lobes (Figs. 239, 240).
PJiiloponcUa tingena males are smaller
than P. .si^natella males, having a cara-
pace length of less than 1.2 mm and a
sternum length of 0.6 mm. Other features
distinguishing these two species are
found in P. si<i,n(itell(i diagnosis.

Description. Female. Total length
2.8 to 4.0 mm (X - 3.45, SD = 0.30, N =
40), carapace length 1.0 to 1.3 mm (X =

1.14, SD = 0.08), sternum length 0.6 to
0.8 mm (X = 0.74, SD = 0.06). Abdomen
two-thirds as wide as long with two dis-
tinct tubercles in anterior third. Some
specimens entirely black with light gray
rings in center of femora and tibiae of
legs. Lighter species have a dark gray car-
apace with a wide, median, posterior, tri-
lobed white area extending forward 0.6
the carapace length (Figs. 226, 227). The
three lobes are of nearly equal length.
Sternum light reddish brown with gray
posterior lateral margins. Abdominal dor-
sum with three white chevrons or with
closely spaced white pigment spots (Fig.
226, 227). Posterior tip gray. Abdominal
venter with median light gray area.
Legs tan or with color markings as in
darker specimens. Epigynal atrium bor-
dered posteriorly by two lobes which are
widely separated by a shallow depres-
sion (Figs. 230, 231). Atrium open pos-
terolaterally. Distance from posterior
lobes to the anterior rim one-third epig-
ynum's width and five times length of an-
terior rim. In ventral view epigynal open-
ings usually hidden (Fig. 230), but
sometimes visible just anterior and me-
dial to posterior lobes. At their closest
point ducts are separated by a distance of
one diameter.

Male. Total length 2.2 to 2.8 mm (X =
2.50, SD = 0.13, N = 40), carapace length
1.0 to 1.2 mm (X = 1.06, SD = 0.05), ster-
num length 0.6 mm (X = 0.60, SD =
0.02). Carapace light tan often with light
gray sides (Fig. 224). Sternum light tan.
Dorsum of abdomen light tan with gray
posterior tip and a few scattered white
pigment spots (Fig. 224). Venter light tan
or with a very faint gray median area.
Legs light tan with a distal gray ring
around tibiae and metatarsi of some or
all legs. Semicircular median apophysis
palpal bulb with an average diameter of
0.2 to 0.3 mm (X = 0.26, SD = 0.02, N =
40) (Fig. 232).

Species Variability. In Panama and
Colombia P. tingena is sympatric with P.
republicana (Map 3). In the Colombian

Revision of Uloboridae • Opell 529

Plate 8. A. Web of Philoponella divisa female holotype. B. Philoponella divisa female holotype (arrow) flanging from
a signal line at the orb's perimeter. C. Philoponella divisa female (arrow) feeding wfiile flanging from a signal line at tfie
orb's perimeter. D. Web of Philoponella tingena from El Queremal, Colombia (f^^ap 3) witfi three signal lines extending
through a vacant web sector (F). E. Philoponella tingena web from El Queremal, Colombia (Map 3) with a single signal
line. F. Close-up of triple signal lines of web shown in D.

530

Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

department of Valle del Cavica three local
populations of Philoponella having indi-
viduals morphologically indistinct from,
but larger than Panamanian P. tin^ena
were studied: one from the faces of large
boulders and roadside banks at El Que-
remal (1600 m), one from a bamboo forest
south of Cali (1000 m), and one from a
lowland deciduous forest near Cali (1000
m). The latter was found in colonies
among buttress tree roots and its individ-
uals produced a nearly horizontal, com-
plete orb-web and rested at the web's
hub while waiting for prey. Individuals
of the former two populations were less
often gregarious and the few colonies ob-
served were small. Members of these two
populations constructed diagonal or ver-
tical webs with a vacant sector through
which one to three signal lines ran (Plate
8-D-F) and rested at the attachment of
the signal line(s) to a peripheral support.
An insufficient web sample was obtained
to permit detailed analysis of the decid-
uous forest population, but the webs of
bamboo forest {P. tin^ena Cb) and El
Queremal (F. tingena B) populations do
not significantly differ (Table 5). Speci-
mens from the bamboo forest are light tan
to white in color with very few dark
markings; whereas, those from the other
two populations are predominantly dark
gray to black. Additionally, the average
egg diameter (for each species ten eggs
from each of three eggsacs were mea-
sured, SD for each species = 0.02 mm)
for Panamanian P. ting,ena was 0.53 mm
and for the El Queremal populations 0.70
mm. Although this evidence suggests
that two or perhaps all of these popula-
tions may be sibling species of P. tin-
gena, additional study is required before
this can be determined. Previous evi-
dence can be challenged by arguments
that: 1. color differences between popu-
lations are no greater than those found
within some Philoponella species, 2.
careful study of intraspecific web vari-
ability is needed to show that web struc-
ture is not influenced by environmental

and predatory pressures, 3. size differ-
ences do not clearly separated the pop-
ulations from one another (Table 4-C, D)
and the trend for Colombian specimens
to be larger than Panamanian specimens
may represent clinal variation, altitudinal
influence, or character displacement, and
4. data have been obtained from only re-
stricted localities.

Distribution. Costa Rica, Panama,
and Colombia (Map 3).

Philoponella signatella (Roewer),
new combination
Table 4; Figures 234-240; Map 3

Uloborus signcitus O. Pickard-Cambridge, 1898,
Biol. Cent.-Amer., Arachnida-Araneida, 1: 264.
One male and one female syntype from Teapa in
the Mexican state of Tabasco, collected by H. H.
Smith, in the British Museum (Natural History),
examined.

Uloborus signatellus Roewer, 1951, Abh. Naturw.
Ver. Bremen, 32(2): 1344. New name for Ulobo-
rus signatus O. Pickard-Cambridge, 1898, op.
cit., preoccupied by Uloborus signatus O. Pick-
ard-Cambridge, 1876, Proc. Zool. Soc. Lond.,
1876: 579 [=U. plumipes Lucas, 1846].

Diagnosis. Along with P. tingena, P.
sigjiatella females are characterized by
having a deep, undivided atrium with
posteriorly directed troughs, posterior
lobes which are as high as wide, an
atrium eight times wider than long, and
openings not visible in ventral view
(Figs. 230, 231, 239, 240). Philoponella
signatella has closely spaced posterior
lobes separated by a deep notch (Figs.
239, 240) rather than widely spaced lobes
separated by a shallow depression as in
P. tingena (Figs. 230, 231). Females of P.
signatella are larger than those of P. tin-
gena, having a total length greater than
4.0 mm and a sternum longer than 0.9
mm (Table 4). Anterior epigynal rim
about half as long as the epigynal atrium
in P. .signatella (Fig. 239), a fifth as long
in P. tingena (Fig. 230). Males of P. sig-
natella and P. tingena are distinguished
from others of their species group by the
median apophysis bulb's lack of a prom-
inent lobe which, when viewed apically.

Revision of Uloboridae • Opell 531

Table 5. Web characters of Philoponella divisa from the Anchicaya Valley of Colombia (Map

3), P. TINGENA-B from EL QUEREMAL, COLOMBIA (MAP 3), P. TINGENA-CB FROM A BAMBOO PATCH NEAR

Cali, Colombia (Map 3), and P. republican a from the Anchicaya Valley of Colombia (Map 3).

No.

No. SPIRALS

DIAMETER
FREE ZONE

DIAMETER
CENTER MESH

AVERAGE
SPIRAL SPACING

FRAMEWORK

RADII

MAX.

MIN.

MAX.

MIN.

MAX.

MIN.

DOVJN

SIDE

WIDTH

LENGTH

P. divisa

19-31

L2-33

7-16

(mm)
26-48

(mm)
23-48

(mm)
10-18

(mm)
7-15

(mm)
2. 3- 3 .7

(mm)
2.9-3.8

(mm)
Z3-3.7

(cm)
9-22

(cm)
10-20

1 singal
line

no vacant
sector

N=13

RANGE

MEAN

22.4

18.6

10.2

34. 6

29.6

13.8

11.1

3.1

3.3

3.2

14.5

14.2

S.D.

3.0

6.1

2.7

6.9

7.0

2.3

2.0

0.5

0.3

0.4

3.2

P. tingena - B

20-24

15-27

7-12

46-96

41-74

15-24

13-18

4.1-5.6

55-8.8

5.1-8.3

15-34

24-45

1-3 signal
lines

vacant

sector

N=6

RANGE

MEAN

22.3

19.7

9. 5

65.4

55.6

18.9

15.0

4.9

7.2

6.6

21.8

34.9

S.D.

1.4

4. 5

2.2

18.7

13.4

3.2

2.2

1.1

1.2

1.5

7.1

9.8

P. tingena -Cb

19-22

20-29

8-10

43-77

39-60

15-24

12-18

35-6.8

4.0-4.8

3.5-5.0

11-16

18-21

1-2 signal
lines

vacant

sector

N = 4

RANGE

MEAN

20.5

23.8

9.0

59.9

49.7

18.1

14.8

4.5

4. 3

4.1

13.9

19.8

S.D.

1.3

4.1

0.8

12.5

8.4

4.0

2.5

1.6

0.4

0.7

1.9

1.7

P. republicana-

50.4

44. 5

21.2

19.7

2.9

4.0

4.3

18.2

22.1

1 signal line
no vacant sector

extends along the median surface of the
spur's basal region (Fig. 203). Philopo-
nella signatella males are larger than
those of P. ting^ena, having a carapace
length of 1.2 mm or more and a sternum
length of 0.7 mm or more. The median
apophysis bulb of P. signatella is large
and somewhat rectangular (Fig. 235);
whereas, that of P. tingena is smaller and
semicircular (Fig. 232).

Description. Female. Total length
4.0 to 6.7 mm (X = 5.08, SD = 0.69, N =
29), carapace length 1.2 to 1.8 mm (X =
1.59, SD = 0.12), sternum length 0.9 to
1.2 mm (X = 1.07, SD = 0.07). Abdomen
about twice as long as wide with two low
adjacent tubercles in anterior third of
dorsum (Fig. 234). Color similar to U. tin-
gens, but in dark forms median light area
of carapace about half as long as carapace.

In light forms this white area extends an-
teriorly three-quarters the carapace length
and the median lobe is twice as long as
lateral lobes. Dorsal abdominal pattern as
shown in Fig. 234. Epigynal crypt opens
posteriorly and has two closely spaced
posterior lobes separated by a deep
depression (Fig. 239). Atrium's length
about one-fifth the epigynum's width and
two times the width of the anterior rim.
In ventral view epigynal openings ob-
scured by posterior lobes (Fig. 239).

Male. Total length 2.8 to 3.6 mm (X =
2.30, SD = 0.33, N = 4), carapace length
1.2 to 1.4 mm (X = 1.24, SD = 0.09), ster-
num length 0.7 to 0.8 mm (X = 0.75, SD =
0.04). Carapace light tan with wide, di-
verging lateral gray lines (Figs. 237).
Sternum light tan with gray posterolater-
al margins. Abdomen's dorsum with nu-

532 Bulletin Museum uf Comparative Zoology, Vol. 148, No. 10

meroiis white spots and gray posterior
tip. Venter with faint gray median area
bordered by a few white spots. Legs tan
or with faint gray distal rings on tibiae
and metatarsi. Median apophysis bulb
of the palpus has an average diameter of
0.4 to 0.5 mm (X = 0.43, SD = 0.03, N =
4) and is rounded to reetangular in shape
(Fig. 235).

Distribution. Honduras through east-
ern Mexico (Map 3).

Philoponella vicina (0. Pickard-Cam-
bridge), new combination
Figures 241-244

Ulobonis vicinus O. Pickard-Camhridjie, 1898,
Biol. Cent.-Amer., Arachnida-Araneidea, I, 266.
Female holotype from Teapa in the Mexican state
of Tabasco, collected by H. H. Smith, in the Brit-
ish Museum (Natural Histor\), examined.

Note. This species is known only
from the female syntype, upon which the
following description is based.

Diagnosis. Females of P. vicina can
be distinguished from those of P. tingena
and P. signatella by having posterior
epigynal lobes which are wider than
long, by having laterally rather than pos-
teriorly directed epigynal troughs, by
having openings visible in ventral view,
and by having an epigynal atrium which
is two-thirds rather than one-third the
length of the epigynum (Figs. 242, 243).
PJiiloponella vicina is distinguished from
P. repiiblicana by having the epigynal
openings in depressions at the center of
the epigynum rather than on raised areas
in the posterior third, by having an an-
terior rim one-eighth rather than one-six-
teenth the length of the epigynum, by
having a total body length less than 4.2
mm, and by having a tan rather than black
sternum which is less than 0.8 mm long.

Description. Female. Total length
4.2 mm, carapace length 1.2 mm, sternum
length 0.8 mm. Abdomen slightly more
than half as wide as long with two small,
but distinct humps in anterior quarter
(Fig. 241). Carapace dark gray with a
wide median light area extending ante-
riorly almost to the posterior eye row.
Sternum tan with gray posterior lateral
margins. Dorsum of abdomen with broad,
light median area broken only by a gray
patch between the humps and a smaller
gray patch midway along the abdomen's
length (Fig. 241). Sides of abdomen light
gray, posterior tip dark gray. Venter of
abdomen with a light gray median area.
First femur with a distal light ring, sec-
ond through fourth femora and all tibiae
with proximal and median light rings.
Metatarsi with a distal gray ring. Epig-
ynal crypt opens laterally and is half as
long as wide. Each opening is in a long
depression at the anterolateral border of
a wide posterior lobe (Fig. 242, 243). An-
terior rim about one-fifth the atrium's
length. Ducts separated by five of their
diameters, each leading first to a bulb
with a diameter half that of the sperma-
theca and then looping before connecting
to a spherical spermatheca (Fig. 244).

Distribution. Southern Mexico.

The Philoponella semiplumosa
Species Group

Philoponella semiplumosa, P. oweni, P.
arizonica, and P. vittata belong to this
species group. Clypeus height of females
eciual to AME diameter. Clypeus height
of males 1.5 to two AME diameters. Epig-
ynal openings near the center of the
atrium and separated by a distance equal
to at least 0.8 the atrium's width (Figs.

Figures 235-240. Philoponella signatella (Roewer). 235. Retrolateral view of male left palpus. 236. Retrolateral view of
expanded male left palpus. 237. Dorsal view of male. 238. Lateral view of epigynum. 239. Ventral view of epigynum. 240.
Posterior view of epigynum. Figures 241-244. Philoponella vicina (O. Pickard-Cambridge). 241. Dorsal view of female.
242. Ventral view of epigynum 243. Posterior view of epigynum. 244. Dorsal view of cleared epigynum. Figure 245. Apical
views of left median apophysis spurs of male palpi of ttie Philoponella semiplumosa species group. Figures 246-247.
Philoponella semiplumosa (Simon) dorsal views of females.

Revision of Uloboridae • Opell 533

534 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

P. arizonica \\ /-^ Af

o P. oweni

• P. semiplumosa

Map 4. Distribution of Philoponella arizonica, P. oweni, and P. semiplumosa.

250, 259, 266). Anterior and posterior atri-
al rims contiguous and lateral troughs ab-
sent. Anterior rim narrow and precipitous
and posterior rim continuous without
clearly defined lateral lobes (Figs. 251,
262, 267). Except in P. vittata (Fig. 268),
each duct spirals tightly around itself be-
fore connecting with a spherical sper-
matheca (Fig. 252). Male palpus' conduc-
tor blade elongate and nearly three times
as long as wide (Figs. 253, 270). Conduc-
tor spike longer than the broad median
apophysis spur's apical region (Figs. 25.3,
270). Median apophysis bulb encompas-
ses the median apophysis spur's basal re-
gion along 0.7 of its perimeter (Fig. 245).
Retrolateral palpal femoral tubercle lon-
ger than prolateral tubercle and its length
about equal to its basal width (Fig. 269).

Philoponella semiplumosa (Simon),
new combination
Table 6; Figures 245-254; Map 4

Uloborus semiplumosus Simon, 1893, Ann. Soc.
ent. Fr., 61: 424. Three females from Caracas and
San Esteban, Venezuela collected 27 December
1887 to 20 January 1888 and 29 February to 27
March 1888, respectively by E. Simon, in Mu-
seum National d'Histoire Naturelle, Paris, ex-
amined.

Uloborus variegatus O. Pickard-Cambridge, 1898,
Biol. Cent.-Amer., Arachnida-Araneidea, I: 266.
Four female and four immature syntypes from
Omilteme(i) in the Mexican state of Guerrero,
collected by H. H. Smith, in the British Museum
(Natural History), examined, NEW SYNONYMY.

Uloborus ahstrusus Gertsch and Davis, 1942, Amer.
Mus. Novitates, 1158: 18. Male holotype and fe-
male allotype from Xilitla in the Mexican state of
San Luis Potosi, collected 22 December 1939, fe-
male and four immature paratypes from the same

Revision of Uloboridae • Opell 535

Table 6. Analysis of species in the Philoponella semiplumosa species group.

0.8

0.7

0.6-

0.5-

1
fi

0.4-

DqS^

□^^D^

0.3-
0.2-

/^ 1

— 1 1 —

— 1

0.5 0.6 0.7

qOo

•• •

n P. semiplumosa
• P. oweni
O P. arizonica

0.8 0.9 1.0 1.1

sternum length

1.2 1.3 1.4 1.5

0.8

0.7

0.6

0.5

d
t

0.4

0.3

0.2

DOT

•• • ^ •

.:s.::A::

ti@«Mfa

□ D n i&n
H nnnnng

• D •J ••

5§go
o

o o
o

n P. semiplumosa
• P oweni
O P- arizonica

-^/ — h

0.7 0.8 0.9 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7

sternum length

536 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

locality, collected 2 December 1939 by A. M. and
L. J. Davis, in the American Museum ot Natural
History, examined, NEW SYNONYMY.

Diagnosis. Females are similar to
those ot P. owcni and P. arizonica, hut
smaller, having a total length of less than
4.6 mm, a carapace length of less than 1.5
mm, and a sternum length of less than 1.0
mm. In females of P. scmipJumosa the
anterior epigynal rim about as long as the
posterior rim (Fig. 250), rather than one-
fourth as long (Figs. 257, 259). Internal
epigynal ducts usually do not show
through the atrium's integiunent as they
do in the other two species. Venter of fe-
male's abdomen with narrow (Fig. 249)
rather than wide (Fig. 256) paraxial white
stripes setting off a median longitudinal
gray stripe. Males are similar to those of
P. oweni and P. arizonica but are small-
er, having a carapace length of less than
1.4 mm, a sternum length of less than 0.8
mm, and a bulb width of less than 0.4 mm.

Description. _ Female. Total length
3.6 to 4.6 mm (X = 3.94, SD = 0.25, N =
37), carapace length 1.1 to 1.5 mm (X =
1.25, SD = 0.11), sternum length 0.7 to
1.0 mm (X = 0.85, SD = 0.07). Carapace
dark gray with light, trilobed white patch
in posterior half or black without light
markings (Figs. 246-247). Sternum gray
to black. Al)domen with prominent dorsal
humps in anterior third. Dorsum light
gray with white median area and dark
gray sides (Fig. 246) to black with a pos-
terior white chevron and three pairs of
white spots (Fig. 247). Venter of abdo-
men gray to black with narrow paraxial
stripes (Fig. 249). Legs brown to gray.
First femur with light distal ring, second
through fourth femora with light proxi-
mal and distal rings (proximal ring may
be lacking in darker specimens). Tibiae
with light proximal and median rings.

First through third metatarsi and tarsi
with narrow, light proximal ring, fourth
metatarsus with narrow, light proximal
and distal rings. Epigvnal atriiun 0.3 to
0.4 mm wide (X = 0.36, SD = 0.05, N =
55) and three-fifths as long as wide. An-
terior rim of epigynum about as wide as
posterior rim (Fig. 250). Epigynum in
posterior view with a median depression
(Fig. 251).

Male. Total length 2.4 to 3.4 mm (X =
2.93, SD = 0.35, N = 25), carapace length
1.0 to 1.4 mm (X = 1.21, SD = 0.10, N =
30), sternum length 0.6 to 0.8 mm (X =
0.70, SD = 0.05, N = 36). Carapace tan
with a light median stripe beside which
are brown, diverging stripes (Fig. 248).
Sternum tan dusted with gray to solid
gray in darker specimens. Dorsum of ab-
domen white with a pair of dark dots in
the anterior third (Fig. 248) or, in darker
specimens, with three to four dark gray
chevrons. Venter of al^domen with a
slightly darker median area. Legs tan,
first femur reddish brown. The first tibia
of most with eight or fewer dorsal spines.
Length of central projection of median
apophysis about 1.5 times the width of its
base (Fig. 253). Bulb 0.3 to 0.4 mm wide
(X = 0.37, SD = 0.05, N = 36).

Distribution. SE Texas through Cen-
tral American and the Caribbean Islands
(Map 4).

Philoponella oweni (Chamberlin),
new combination
Table 6; Figures 255-258; Map 4

Ulohorus oweni Chamberlin, 1924, Proc. Calif.
Acad. Sci., 12(28): 579. Female holotype from
Gulf of California, Carmen Island, Marquer Bay,
collected 23 May 1921 by J. C. Chamberlin, V.
Owen, and 1. M. Johnston, in the California Acad-
emy of Sciences (specimen No. 1363), examined.

Figures 248-254. Philoponella semiplumosa (Simon). 248. Dorsal view of male. 249. Ventral view of female abdomen.
250. Ventral view of epigynum. 251. Posterior view of epigynum. 252. Dorsal view of cleared epigynum. 253. Retrolateral
view of left male palpus. 254. Eggsacs. Figures 255-258. Philoponella oweni (Cfiamberlin). 255. Dorsal view of female.
256. Ventral view of female abdomen. 257. Ventral view of epigynum. 258. Dorsal view of male. Figures 259-261.
Philoponella arizonica (Gertsch). 259. Ventral view of epigynum. 260. Dorsal view of female. 261. Dorsal view of male!

Revision of Uloboridae • Opell 531

538

Bulletin Museum of Coin punitive Zoology, Vol. 148, No. 10

Muma and Cieitsch, 1964, Amer. Mus. Novitates
2196: 34.

Diagnosis. Females are similar to
those of P. scmiplumosa and P. arizoni-
ca, but are intermediate in size, having
a carapace length of 1.5 to 2.0 mm and a
sternum length of 1.0 to 1.5 mm. Like P.
arizonica and unlike P. scmiplumosa, the
anterior epigynal rim is one-fourth as
long as, rather than e(iual in length to, the
posterior rim; some of the internal ducts
show through the atrium's integument
(Fig. 257), and broad, paraxial stripes set
off a median gray area on the venter of
the abdomen (Fig. 256). Females of P.
oiveni differ from those of P. arizonica by
being more darkh' pigmented, by having
an epigynal atrium which is three rather
than two times as wide as long (Fig. 257),
and by lacking ventral lateral dark lines
(Fig. 262) on the posterior face of the
epigynum. Males of P. owcni are similar
to those of P. scmiplumosa and P. ari-
zonica but are intermediate in size, hav-
ing a carapace length of 1.6 to 1.8 nmi, a
sternum length of 0.9 to 1.1 mm, and a
bulb width of 0.5 to 0.6 mm. Pliiloponclla
owcni has a median apophysis spine the
length of whose central projection is 1.5
the width of its base rather than 2.5 as in
P. arizonica (Fig. 245).

Description. _ Female. Total length
4.6 to 7.1 mm (X = 5.81, SD = 0.69, N =
27), carapace length 1.5 to 2.0 mm (X =
1.82, SD = 0.17, N = 27), sternum length
1.0 to 1.5 mm (X = 1.21, SD = 0.10, N =
46). Carapace brown with broad, median
light area extending anteriorly to eyes
(Fig. 255). Sternum tan at center with
gray margins. Abdomen with small
humps about midway along its length,
dorsum entirely white or with white me-
dian area and gray sides (Fig. 255). Ven-
ter of abdomen white with narrow gray
median longitudinal stripe (Fig. 256).
Legs light brown. First femur with distal
light ring, second through fourth femora
with proximal and distal light rings. Tib-
iae with proximal and median light rings.
First metatarsus without rings, second

through fourth metatarsi with faint
proximal and distal light rings. Tarsi
with light narrow proximal rings. Epigy-
nal atrium 0.4 to 0.5 mm wide (X = 0.46,
SD = 0.05, N = 46) and about two-fifths
as long as wide. Anterior rim of epigy-
num one-fourth the width of posterior
rim (Fig. 257). Epigynum in posterior
view without a proiment median depres-
sion.

Male. Total length 3.2 to 4.4 pm (X =

4.00, SD = 0.33, N = 14), carapace length
1.6 to 1.8 mm (X = 1.69, SD = 0.09, N =
14), sternum length 0.9 to 1.1 mm (X =

1.01, SD = 0.07, N = 30). Carapace brown
dusted with gray, light median longitu-
dinal stripe extends anteriorly to eyes
(Fig. 258). Sternum with gray margins
and tan center. Dorsum of abdomen all
white or with a white median area and
gray sides (Fig. 258). Venter of abdomen
light or with a narrow median gray area
set off by broad paraxial white stripes.
Legs reddish brown and, in most speci-
mens, with no prominent markings. First
tibia of most with nine or ten dorsal mac-
rosetae. Length of central projection of
median apophysis about 1.5 the width of
its base. Bulb 0.5 to 0.6 mm wide (X =
0.52, SD = 0.03, N = 30).

Distribution. SW Texas, SW United
States, and NW Mexico (Map 4).

Phitononella arizonica (Gertsch),
new combination
Table 6; Figures 259-263; Map 4

VIohonis arizonicus Gertsch, 1936, Amer. Mus.
Novitates, 852: 2. Male holotype from Sal)ino Ba-
sin, Santa Catalina Mountains, Arizona (elevation
1280 m), collected 8 to 12 July 1916 by F. E. Lutz,
in the American Museum of Natural History, ex-
amined. Muma and Gertsch, 1964, Amer. Mus.
Novitates, 2196: 35.

D/c/gnosis. Males and females are
similar to those of P. oweni but are usu-
ally larger, males being at least 4.0 mm
long and females being at least 6.7 mm.
Females of P. arizonica are more light
colored. See P. owcni diagnosis for ad-

Revision of Uloboridae • OpelJ 539

ditional distinguishing characters of these
two species.

Description. Female. Total length
6.7 to 8.0 mm (X = 7.26, SD = 0.50, N =
11), carapace length 2.3 to 3.2 mm (X =
2.48, SD = 0.21, N = 16), sternum length
1.5 to 1.7 mm (X = 1.52, SD = 0.05, N =
16). Carapace brown with broad, light
median longitudinal stripe extending an-
teriorly to eyes (Fig. 260). Sternum tan,
often dusted with gray. Abdomen with
small humps in anterior two-fifths, dor-
sum all white or with light gray sides
(Fig. 260). Venter of abdomen all white
or with a narrow median gray stripe. Legs
light tan with no prominent markings.
Epigynal atrium 0.6 to 0.8 mm wide (X =
0.68,' SD = 0.04, N = 16), and four-sev-
enths as long as wide (Fig. 259). Anterior
rim of epigynum one-fourth the width of
the posterior rim. Epigynum in posterior
view without a prominent median
depression and with ventral lateral dark
lines showing through the integument
(Fig. 262).

Male. Total length 4.0 to 6.4 mm (X =
5.34, SD = 1.03, N = 11), carapace length
2.0 to 3.8 mm (X = 2.56, SD = 0.50, N =
11), sternum length 1.1 to 1.4 mm (X =
1.27, SD = 0.12, N = 11). Carapace tan
with two brown paraxial stripes extend-
ing to anterior margin (Fig. 261). Sternum
tan, in some dusted with gray. Dorsum of
abdomen all white or with gray sides and
cardiac area (Fig. 261). Venter of abdo-
men all white or with narrow gray me-
dian longitudinal stripe. Legs tan with no
prominent markings. First tibia of most
with 12 dorsal macrosetae. Length of cen-
tral projection of median apophysis about
2.5 times the width of its base. Bulb of
median apophvsis 0.6 to 0.7 mm wide
(X = 0.66, SD = 0.04, N = 10).
Distribution. Arizona (Map 4).

Philoponella vittata (Keyserling),
new combination
Figures 264-271; Map 5

Uloborus vittatus Keyserling, 1882, Verb. zool.-l)ot.
Ges. Wien, 31: 279. Three female and one male

syntypes from Peioi, in British Museum (Natural
History), examined.

Uloborus serviilus Simon, 1892 [1893], Ann. Soc.
ent. France, 61: 424. Male, three female, and
three immature female syntypes from San Este-
ban (6 km S of Puerto Cabello) in the Venezuelan
state of Caracas, collected 29 February to 27
March by E. Simon, in Museum National
d'Histoire Naturelle, Paris, examined, NEW
SYNONYMY.

Uloborus scmiar<i.enteu.s Simon, 1893, Ann. Soc.
ent. France, 62: 299. Six female, two male, and
fi\e immature syntypes from le Para (presently
Belem), Brazil, collected by M. de Mathan, in
Museum National d'Histoire Naturelle, Paris, ex-
amined, NEW SYNONYMY.

Uloborus (n7}azonicus Mello-Leitao, 1949, Boletim
do Museu Nacional, Rio de Janeiro, 92: 2. Female
holotype from the vicinity of Rio Xingu in north-
western Brazil, collected by J. C. Car\alho, in the
Museu Nacional, Rio de Janeiro, examined, NEW
SYNONYMY.

Note. Keyserling lists female and
male syntypes from "Peru: Amable Maria
and Junin" and mentions that numerous
examples are in the University ol Warsaw
collection. These specimens were not re-
ceived with other requested material
from the Polska Akademia Nauk and ap-
pear to be lost. Specimens in the British
Museum (Natural Histoiy) marked "type"
appear to be part of Keyserling's type se-
ries.

Diagnosis. Females are most similar
to those of P. republicana and P. vicina,
but may be distinguished by their color
pattern (Fig. 264), by having a broad, con-
vex anterior epigynal border (Fig. 266)
rather than a narrow, precipitous margin;
by having openings in the posterior sixth
rather than posterior third or half of the
atrium (Fig. 266), and by having these
openings separated by a third the atrial
width. Epigynal ducts loop twice before
connecting with the spermathecae (Fig.
268). Males are similar to those of the oth-
er members of this species group, but
may be distinguished by the presence of
a light chevron on the abdomen's venter
(Fig. 265), by having a median apophysis
bulb which is dome-shaped rather than
flat in lateral view, and by having a more
extensive conductor lobe (Fig. 270).

Description. Female. Total length

540 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

4.7 to 6.3 mill (X = 5.29, SD = 0.46, N = ameter. Sternum light tan. Alxlomen's

16), carapace length 1.4 to 1.7 mm (X = dorsum light tan, often with light gray

1.58, SD = 0.10), sternum length 0.9 to sides, four medially divided white chev-

1.1 m (X = 1.02, SD = 0.07). Carapace ions, and two anterolateral white patches

uniformly tan to dark brown or with a (Fig. 265). Abdominal venter with a gray

light central area which may be trilobed median longitudinal stripe about half the

(Fig. 264). The AMEs often surrounded abdomen's width and with a light chev-

by a small, light area, never by a black ron midway between the spinnerets and

area. Clypeus height 0.5 to 0.6 AME di- epigastric furrow. Legs all light or faintly

ameter. Sternum brown to dark gray. Ab- colored as in females. Palpal femur with

domen half as wide as long with a slight two small, equal-sized proximal ventral

raised area in the anterior quarter. Dor- tubercles. Median apophysis bulb large

sum white to dark gray with a light me- and dome-shaped, conductor spike long

dian longitudinal stripe a third to half the and conductor blade large, three times as

abdomen's width, within which are three long as wide (Fig. 270).

pairs of darkly bordered white spots or Distribution. South America (Map 5).
three medially divided chevrons (Fig.

264). Anterior lateral margins of abdomen ^^^ Rhiloponelia fasciata

each wi h a white oval patch, posterior 3 .^^ ^
tip or al)domen with a white clievron.

Abdomen's venter with an hourglass- This group contains P. fasciata, P.
shaped median light area set off by a nar- para, and P. bella. Females of these
row white line. Booklung covers unpig- species and males of F. para have a clyp-
mented. Legs uniformly light in color or eus height equal to the AME diameter
with dark proximal and distal rings of and male P. fasciata a clypeus height
femora and tibiae. Epigynum with a shal- twice the AME diameter. Epigynal open-
low central depression bordered ante- ings are found in the anterior third (Fig.
riorly and laterally by broad, convex 287) or posterior half of the atrium (Fig.
margins and posteriorly by a narrow, 275, 284) and are separated by a distance
overhanging rim (Fig. 266). Openings in equal to 0.6 to 0.8 the atrial width. The
posterior sixth of epigynum, separated by duct leading from each opening coils
a third the epigynal width. Ducts loop once (Figs. 275, 285) before connecting
twice before entering spherical sperma- to the spermatheca and in the region of
theca (Fig. 268). this coil its walls are thickened. Anterior
Male. Total length 3.6 to 4.2 mm (X = epigynal rim precipitous, its anterior bor-
3.91, SD = 0.19, N = 16), carapace length der is broad and convex (Figs. 275, 284,
1.5 to 1.8 mm (X = 1.63, SD = 0.09, ster- 287). Anterior and posterior rims are not
num length 0.9 to 1.0 mm (X = 0.96, SD = separated by conspicuous troughs and
0.04). Carapace light tan, often with a the posterior rim is not indented (Figs.
light median stripe and lateral markings 276, 286, 288). The male palpus has a
(Fig. 265). All eyes with a narrow black short, square conductor blade, but unlike
rim. Clypeus height ccjual to AME di- the other species groups, has a long, thin.

Figures 262-263. Philoponella arizonica (Gertsch). 262. Posterior view of epigynum. 263. Dorsal view of cleared epig-
ynum Figures 264-271 Philoponella vittata (Keyserling). 264. Dorsal view of female. 265. Dorsal view of male. 266.
Ventral view of epigynum. 267. Posterior view of epigynum. 268. Dorsal view of cleared epigynum. 269. Ventral view of
male palpal femur 270 Retrolateral view of male left palpus. 271. Apical view of left male palpus. Figure 272. Apical
views of male left median apophysis spurs of Philoponella fasciata species group. Figures 273-274. Philoponella fasciata
(Mello-Leitao). 273. Female carapace. 274. Dorsal view of female holotype abdomen.

Revision of Uloboridae • Opell 541

p. fasciata

542

Bulletin Museum of Cuinparative Zoology, Vol. 148, No. 10

• Philoponella vlttata
® Philoponella fasciata

Map. 5. Distribution of Philoponella vittata and P. fas-
ciata.

distal prong which runs nearly parallel to
the conductor spike (Figs. 280, 283). The
conductor spike is as long as or longer
than the median apophysis spur's slender
apical region.

Philoponella fasciata (Mello-Leitao),
new combination

Figures 272-281; Map 5

Ulohoru.s fa.sciatu.s Mello-Leitao, 1917,' Esc. Sup.
AKric. Medic. Vet., 1(1): 4. Female holotype from
Nova-It,niacv'i in the Brazilian state of Rio de Ja-
neiro, collected by Blanc de Freitas, in Museu
Nacional, Rio de Janeiro, examined.

Diagnosis. Females are distinguish-
ed from other members of the variegata
species group by having: 1. a total length
of less than 3.5 mm, 2. a sternum length
of 0.7 mm or less, 3. long, longitudinal
epigynal openings located in the poste-
rior half of the epigynal crypt and bor-
dered laterally by a broad, flat inclined
region (Fig. 275), and 4. a gray spot just
anterior to the carapace's thoracic depres-
sion. Males are distinguished from others
of the genus by their high, nearly cylin-
drical carapace (Figs. 278, 279), long lat-
eral palpal femoral tubercle (Fig. 281),
and coloration (Figs. 278, 279). They are
distinguished from other members of the
variegata species group by their short
basal conductor lobe and its long, thin
extension (Fig. 280).

Description. _ Female. Total length
2.4 to 3.2 mm (X = 2.76, SD = 0.30, N =
10), carapace length 1.0 to 1.1 mm (X =
1.00, SD = 0.05), sternum length 0.6 to
0.7 mm (X = 0.64, SD = 0.04). Abdomen
three-fourths as wide and high as long
and peaked in the anterior third of its
length. Carapace light tan to black, all but
the darkest specimens having color mark-
ings similar to those shown in Fig. 273.
Sternum tan to dark brown. Abdomen
coloration of holotype (Fig. 274) differs
from that of other specimens examined.
In these it ranges from white to black, but
in most specimens has a wide white dor-
somedian stripe, gray sides, and a gray
venter with a pair of thin, widely sepa-
rated paraxial stripes extending from
booklungs to spinnerets. In ventral view
epigynal atrium is 0.1 to 0.2 mm long (X =
0.15, SD = 0.02) and 0.2 to 0.3 mm wide
(X = 0.26, SD = 0.02). Its long openings
lie just posterior to the atrium's center,
medial to wide lateral borders, and sep-

Figures 275-281 Philoponella fasciata (Mello-Leitao). 275. Ventral view of holotype's epigynum. 276. Posterior view of
epigynum 277. Dorsal view of cleared epigynum. 278. Dorsal view of male carapace. 279. Lateral view of male carapace.
280. Retrolateral view of apical region of male left palpus. 281. Retrolateral view of male left palpal femur. Figures 282-
286 Philoponella para n. sp. 282. Ventral view of female abdomen. 283. Retrolateral view of male palpus. 284. Ventral
view of epigynum. 285. Dorsal view of cleared epigynum. 286. Posterior view of epigynum. Figures 287-288. Philopo-
nella bella n. sp. 287. Ventral view of epigynum. 288. Posterior view of epigynum.

Revision of Uloboridae • Opell 543

544 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

arated by a distance e(iual to 0.8 the which protrudes conspicuously ventrally

atrium's width (Fig. 275). In posterior and in posterior view is as high as broad

view posterior phite twice as wide as (Figs. 284, 286). The atrium is twice as

high and with a slight median indenta- wide as long and openings situated in its

tion (Fig. 276). A duct leading from each posterior quarter are hidden in ventral

opening loops once aroimd itself before view (Fig. 284). The abdomen's light

connecting with a spherical spermatheca venter has a dark W-shaped mark (Fig.

(Fig. 277). 282). Males of P. para and P.fasciata are

Male. The male of this species is de- characterized by having a small conduc-
scribed here for the first time. Total tor lobe with a long, thin extension pro-
length 2.2 to 2.6 mm, carapace length 1.1 jecting nearly parallel to the conductor
to 1.3 mm, sternum length 0.6 to 0.7 mm. spike (Figs. 280, 283). The median
Carapace high with nearly straight sides, apophysis bulb of P. para is dome-
a deep and wide transverse thoracic shaped and has a small apical lobe (Fig.
groove, and a more prominent AME tu- 283); whereas the bulb of P. fasciata
bercle than present in other Philoponelhi males is flat (Fig. 280).
males (Figs. 278, 279). Length of retro- Description. Female. Total length
lateral palpal femoral tubercle twice the 2.4 to 2.8 mm, carapace length 0.9 to 1.0
width of its base (Fig. 281). First femur mm, sternum length 0.7 mm. Carapace
with two or three prolateral, one retrolat- and sternum mottled gray. Abdomen
eral, one dorsal, and no ventral macro- width and height two-thirds its length,
setae. First tibial prolateral, retrolateral, Abdominal dorsum white with irregular
and dorsal surfaces each with four spines, patches of gray, lateral surface white to
venter without spines. Carapace tan to gray, venter white with a more or less
black and in most specimens with color distinct, upright gray "W" (Fig. 282).
markings similar to those shown in Figs. Legs white. First and fourth femora with
278, 279. Sternum tan to gray. Abdomen proximal, central and distal gray rings;
tan with irregular white spots and a gray second and third femora with median and
posterodorsal tip. Median apophysis bull) distal gray rings. First tibia with proximal
very flat (Fig. 280), 0.3 mm long and 0.2 and distal gray rings; first metatarsus and
to 0.3 mm wide. Conductor spur long and tarsus gray. Second through fourth tibiae
concave (Fig. 280). Conductor basal lobe and metatarsi with median and distal
shorter than other known members of the gray rings. Second through fourth tarsi
variegata species group and with a long, white. In ventral view epigynal atrium
thin, nearly transparent projection which about twice as wide as long (length 0.1
extends parallel to the conductor spur mm, width 0.2 mm) and has a concave,
and is about as long as the latter (Fig. ventrally directed median lobe which is
280). three-fourths as wide as the atrium and

Distribution. Southeastern Brazil and has a straight ventral margin (Fig. 284).

Paraguay (Map 5). The epigynal atrium's anterior rim is

rounded (Fig. 284). In posterior view the

Philoponella para n. sp. epigynum is as high as wide (Fig. 286).

Figures 282-286 An indistinct epigynal opening is located

Types. Female holotype, one male and two female '^*^ ^'^^'^ centrolateral margin of the atrium

paratypes from Taguaraiapa [cannot he located in '^l^Cl the duct leading from it loops once

gazeteteers] in the Paraguay department of Alto before connecting to a small spherical

Parana in the Ainerican Museum of \atural His- spermatheca which lies at the level of the

tory. Ihe specific epithet is an arhitrarv com hi- >.,i.,,..; 4. • i • /r^- to(-\

nation of letters. ' autei or atrial run (Fig. 285).

Male. Total length 2.2 mm, carapace

Diagnosis. Females are distinguish- length 1.0 mm, sternum length 0.6 mm.

ed by having a posterior epigynal margin Carapace and sternum light tan; lateral

Revision of Uloboridae • Opel! 545

regions of carapace lightly mottled with
gray; sternum with narrow gray lateral
borders. Abdomen tan, overlain with
white patches; dorsal tip gray, venter
with two widely separated, paraxial gray
stripes. Legs tan with a faint, distal gray
ring on first femur. First femur with three
prolate ral, one retrolateral, and no dorsal
and ventral macrosetae. First tibia with
three prolateral, four retrolateral, four
dorsal and no ventral spines. Proximal,
ventral surface of palpal femur with two
small, nearly equal-sized tubercles. Me-
dian apophysis bulb dome-shaped, 0.2
mm long, and 0.2 mm wide, and with a
' small apical lobe (Fig. 283). Median
apophysis spur's basal region two-thirds
as long as wide, with a prominent apical
lobe and a spike ec^ual to the latter's
length (Fig. 272). Conductor's basal lobe
(Fig. 283) longer than that of P. fasciata.
Like P. fasciata, the basal conductor lobe
has an extension which runs nearly par-
allel to the conductor spike. In P. para
this extension is about two-thirds the
spike's length.

Distribution. Known only from the
type locality in Paraguay.

Philoponella bella n. sp.
Figures 287-288

Types. Female holotype and female paraHpe from
Rio Domachui trail, elev. 2700 to 3000 m, near
Santa Marta in the Colombian state of Magdalena,
collected 2 February 1973 by J. A. Kochalka, in
the Museum of Comparative Zoology. The spe-
cific epithet is an arbitrary combination of letters.

Diagnosis. Female P. bella are distin-
guished from other members of the genus
by having an epigynal atrium whose
length and width are equal and whose
protruding posterior surface has two lat-
eral grooves (Figs. 287, 288).

Description. Only females are known.
Total length 3.5 to 3.6 mm, carapace
length L3 to L4 mm, sternum length 0.8
mm. Carapace and sternum mottled gray;
carapace with white lateral rim. Abdo-
men tan, mottled with dark gray and with
a pair of large dark dorsal spots in the
anterior quarter and a cluster of five nar-

row, transverse, gray dorsal stripes in the
posterior third. First femur gray with a
distal white ring. Second through fourth
femora white, each with a proximal, cen-
tral, and distal gray ring. Proximal half of
tibiae white with a gray ring, distal half
gray-brown. Metatarsi white with dark
central and distal rings. Tarsi white
proximally and gray distally. In ventral
view length and width of epigynal atrium
are equal; half of the atrium's length pro-
jecting posterior to the epigastric furrow
(Fig. 287). The atrium's posterior region
forms a deep pocket and is bordered by
a thin posterior rim. Openings are situ-
ated slightly anterior to the rim's center
at the atrium's lateral margins. In poste-
rior view the epigynum has two lateral
grooves (Fig. 288). From each epigynal
opening a duct loops once before con-
necting with a small, spherical sperma-
theca from whose posterior median sur-
face a fertilization duct arises.

Distribution. Known only from the
type locality in northeastern Colombia.

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548 Bulletin Museum of Comparative Zoology, Vol. 148, No. 10

INDEX

Valid names are printed in italics. Page numbers
refer to descriptions, starred page numbers to illus-
trations, and page numbers in parentheses to dis-
tribution maps.

abdominalis, Sijbota 457*, 463*, 491*, 494, 497*
abstrusus, Uloborus 534
aegrotus, Uloborus 504
albicans, Ariston 477*, 480
amazonicus, Uloborus 539
ancepts, Uptiotes 485
Ariston (471), 478
aristus. Ariston 479*, 482

arizonica, Philoponella 533*, (534), 537*, 538, 541*
Astavakra 470
ater, Sylvia 494
ater, Uloborus 470

banibusicola, Miagrammopes 490, 491*
bellu, Philoponella 543*, 545
bituberculata, Orinomana 499*, 500
bucki, Uloborus 504
campestrattts, Uloborus 503*, (505), 506
caraibe, Zosis 510
cavatus, Hyptiotes 483*, 485, 487*
cinereus, Uloborus 506
collinus, Uloborus 470
corticeus, Miagrammopes 491*, 492
costalimae, Uloborus 510
ciuninamensis, Uloborus 524
Cyllopodia 485
Darumuliana (471), 515
delectus, Uloborus 512
dissimila, Tangaroa 476
diversus, Uloborus 463*

divisa, Philoponella 517*, 519*, 521*, 526, 529*
domesticus, Uloborus 510
dubius, Uloborus 512

faseiutu, Philoponella 541*, 542, (542), 543*
festivus, Uloborus 502
formosus, Uloborus 470
geniculatus, Zosis 447*, 449*, 459*, 463*, (505),

509*, 510, 511*
gertsehi, Hyptiotes 454*, 463*
gibbosa, Darumuliana 513*, 515
glomosus, Uloborus 459*, 463*, 499*, 501
grammica, Purumitra 507, 509*
Huanacauria 490
Hyptiotes (471), 485
lactescena, Ponella 513*, 516, 517
lamprus, Orinomus 500
lutens, Miagrammopes 489*
latreillei, Uloborus 510
lineata, Veleda 501
manuueata, Phillyra 501
nuina, Orinomana 499*, 501
maniculatus, Uloborus 504
mazolus, Ariston 453*, 477*, 479*, 481
mendozae, Sybota 496, 497*
Miagrammopes (471), 490

minutus, Uloborus 470

Mithras 485

Mumaia 490

mundior, Uloborus 512, 524

niger, Uloborus 470

octonaria, Oetonoba 459*, 512

Octonoba (471), 511*, 512, 513*

Orinomana (471), 498

Orinomus 498

Orithvia 508

osornis, Sybota 457*, 495, 497*

orsinus, Uloborus 502

oweni. Philoponella 533*, (534), 536, 537*

para, Philoponella 541*, 543*, 544

paradoxus, Hyptiotes 483*, 485

penicillatus, Uloborus 447*,449*, 499*, 503*, 504,

(505)
peruvanus, Uloborus 502
Petrunkevitchia 498
Phillvra 501
Philoponella (471), 518
Philoponus 501
plimiipedatus, Uloborus 504
plumipes, Uloborus Lucas 504
plumipes, Uloborus Mello-Leitao 504
Polenecia (471), 482
Ponella (471), 515
productu, Polenecia 479*, 482, 483*
pteropus, Philoponus 501
Purumitra (471), 507
pusilla, Petrunkevitchia 498
Ranguma 490

referena, Siratoba 453*, 487*, 488
republieana, Philoponella 447*, 449*, 459*, 463*,

517*, 519*, 524, (525)
rubiginosa, Sylvia 494
segregatus, Uloborus 503*, 505, (505)
semiargenteus, Uloborus 539
semiplumosa, Philoponella 533*, 534, (534), 537*
servulus, Uloborus 539
sexmucronatus, Uloborus 470

signatella, Pluloponella 517*, 521*, (525), 530, 533*
signatus, Uloborus 530
similis, Sylvia 494
similis, Miagrammopes 490
simus, Miagrammopes 455*, 463*, 489*, 491
sinensis, Uloborus 512
siru, Siratoba 489*, 490
Siratoba (471), 486
spernax, Uloborus 506
Sybota (471), 493
sxbotides, Uloborus 512
Sylvia 493

tahitiensis, Tangaroa 474, 475*
Tangaroa (471), 474
tetramaculatus, Uloborus 470
thuaitesii, Miagrammopes 490
tingena, Philoponella 517*, 521*, (525), 528, 529*
trilineatus, Uloborus 470

Revision of ULOBomoAE • Opell 549

Uloboridae 469
Ulohonts (471), 501
Uptiotes 485
ursinus, Uloborus 512
varians, Uloborus 512
variegatus, Uloborus 534
Veleda 501

veuusta, Petruukevitchia 498
vicina, Philopunella 532, 533*

vittata, Sylvia 494

vittata, Philoponclla 539, 541*, (542)

Waitkera (471), 476

waitkerensis, Waitkera 475*, 476-477*

walckenaerius, Uloborus 501

williauisii, Orithyia 510

yesoensis, Argyrodes 512

Zosis (471), 508

zosis, Uloborus 510

78 0 7

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