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507.73

CaL«8

NUMBER 287
FEBRUARY 10, 1978

RESULTS OF THE LATHROP CENTRAL AFRICAN REPUBLIC

EXPEDITION 1976, ORNITHOLOGY

By Herbert Friedmann

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCENC6

Published by the NATURAL HISTORY MUSEUM
OF LOS ANGELES COUNTY
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CONTRIBUTIONS IN SCIENCE contain articles in the earth and life sciences, presenting
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Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

RESULTS OF THE LATHROP CENTRAL AFRICAN REPUBLIC
EXPEDITION 1976, ORNITHOLOGY1

By Herbert Friedmann2

Abstract: Some 400 specimens of 94 species of birds were collected in the
vicinity of the Ouossi River, in the southern, forested part of the Central African
Republic, by the Lathrop Expedition in June 1976. Forty-three of these species
are here recorded for the first time from that country. Others, known from there
from one or a few earlier specimens, constitute sizeable extensions of previ-
ously known ranges. Aside from these distributional data, weights of all the
birds are recorded as well as the gonadal condition of each specimen, informa-
tion of a sort lacking until now for the birds of that portion of central Africa.

INTRODUCTION

This paper presents the results of the Lathrop Central African Republic
Expedition in ornithology carried out in 1976. All the bird collecting was done
from one base camp, near the Ouossi River, about 11 k west of Baroua,
elevation 680 m, roughly 5°20’N-24°20’E. This is a forested area in the extreme
southern part of what Chapin (1932:90) termed the Ubangi-Shari Savanna Dis-
trict in his map of the faunal areas of Africa. The collecting was done by
Andrew Williams and his two African assistants and skinners, Julius Kyongo
and Philip Imbayi, between 31 May and 22 June 1976. In his report to the
museum, Williams wrote that the forests near the camp site proved to be of
three separate types which he found had small differences in their bird life: 1)
the mature riverine forest along the Ouossi River, with the largest trees of the
three, and, for the most part, dense, lush undergrowth; 2) an area of mature,
but more open forest away from the river north of the camp, somewhat drier,
and with no little streams running through it; 3) a still more open type of
woodland bordering on open grassland. The forests visited were fairly small and
restricted and were some distance west of the larger supposedly isolated,
lowland forest shown in Map B in Hall and Moreau’s Atlas of African Ornithol-
ogy (1970), but, judging from the birds obtained in them, they were similar to, but

Review Committee for this Contribution
Charles T. Collins
Ralph W. Schreiber
Kenneth E. Stager

2Director Emeritus, Natural History Museum of Los Angeles County, 900 Exposition
Boulevard, Los Angeles, California 90007.

Figure 1. Map of Africa showing the Central Africa Republic and the locality of the present collection near
Baroua.

1978

The Lathrop Ornithological
Central African Expedition

3

somewhat less humid than the northern parts of the great Congo forest of Zaire
to the south. During his collecting, involving both mist nets and shotgun,
Williams became aware of the absence of starlings and of akalats ( Sheppardia ) in
all three of these forested areas. It is not conclusive that these birds do not occur
there, as at least 6 species of Lamprotornis and 1 of Sheppardia have been
reported earlier from the Central African Republic, but they cannot be abundant
near the Ouossi River, or they would have been noted.

A collection of 400 specimens of 94 species, made during a little over 3
weeks in the field, cannot be looked upon as a complete representation of a
local, tropical forest bird fauna. However, it is significant in filling gaps in our
knowledge of the distribution of central African birds. The collecting was
confined almost wholly to the forested areas, but a few birds were taken in the
open grasslands immediately adjacent to them. The collector carefully and
conscientiously recorded in detail the colors of the soft parts (eye, bill, bare
skin areas and feet), as well as the weight, the state of the gonads (unmeasured)
and the contents of the stomach of every specimen. The weights and gonadal
condition reported here are the first for their area. The colors of the soft parts
and the stomach contents are included only for species of which Chapin ( 1932-
1954) had little material, and only for some species of special interest has
systematic discussion been thought useful.

In evaluating the extensions of known ranges of a large number of the
included species, the maps of each of the passerines in Hall and Moreau’s atlas
are of the first importance. The maps in Mackworth-Praed and Grant’s two
volumes on the birds of west-central and western Africa show mere black
swatches to indicate ranges, and these are not precise in the sense that are
those of Hall and Moreau, which attempt to show every locality of record. In
all such species maps, where the swatches suggest distributional limits beyond
those given by Hall and Moreau, and where there are no published records in
the earlier literature, I have assumed they were not based on actual specimens
or on definite observations, and have been guided basically by the correspond-
ing maps in Hall and Moreau. Assuming a period of three years from the
completion to the publication of these maps, I have gone back to 1967 in my
search of the literature for possible unmapped records from the Central African
Republic.

Table 1 lists birds in the Lathrop Expedition’s collection which are the
first records in print for their species from the Central African Republic. A
number of others fill sizeable gaps in the specimen records mapped by Hall and
Moreau, but are not new for that political area.

The systematic order and the nomenclature used in this paper is that of
Mackworth-Praed and Grant (1970-1973), the most recent complete work on
the birds of western and central Africa. In a few species it has been deemed
better not to follow their treatment. Further, the generic and specific names
and sequence have been changed, where needed, to conform with the Refer-
ence List of the Birds of the World, by Morony, Bock and Farrand (1975).

4

Contributions in Science

No. 287

TABLE 1

Species of birds hitherto unreported from the Central African Republic.

Pachycoccyx audeberti
Cuculus clamosus clamosus
Cuculus clamosus gabonensis
Cercococcyx mechowi
Alcedo leucogaster leopoldi
Halcyon badia

Halcyon malimbica malimbica
Tockus hartlaubi grand
Pogoniulus scolopaceus flavisquamatus
Pogoniulus bilineatus leucolaema
Pogoniulus subsulphureus flavimentum
Pogoniulus atroflavus
Trachyphonus purpuratus purpuratus
Melignomon zenkeri
Indicator maculatus stictithorax
Indicator exilis exilis
Melichneutes robustus
Campethera nivosa herberti
Smithornis capensis camarunensis
Pitta angolensis longipennis
Pycnonotus gracilirostris chagwensis
Baeopogon indicator indicator

Erythropygia leucosticta collsi
Cossypha natalensis intensa
Alethe diademata woosnami
Alethe poliocephala carruthersi
Trichastoma albipectus albipectus
Trichastoma rufipenne rufipenne
Ptyrticus turdinus

Camaroptera superciliaris flavigularis
Camaroptera chloronota toroensis
Macrosphenus flavicans hypochondriacus
Platysteira castanea castanea
Trochocercus nigromitratus
Anthreptes rectirostris tephrolaema
Nectarinia seimundi traylori
Nectarinia cyanolaema octaviae
Nigrita bicolor brunnescens
Nigrita canicapilla canicapilla
Mandingoa nitidula schlegeli
Spermophaga ruficapilla ruficapilla
Malimbus nitens microrhynchus
Dicrurus atripennis

SPECIES ACCOUNTS

Species here reported for the first time from the Central African Republic
are marked with an asterisk.

Family Accipitridae

Accipiter tachiro canescens (Chapin). — Two adult males and one young,
but fully grown female of this hawk were collected 2, 13 and 16 June. The
subspecies must be commoner in the Ouossi River area than might have been
expected, as it has been a seldom collected bird in most parts of its range. The
female was in worn plumage with a few adult feathers coming in on the upper
wing coverts, and also had a very few pale chestnut feathers showing on the
sides of the otherwise white breast and abdomen. The two males weighed 194
and 205 g, the female 325 g; both adults had enlarged gonads. One of the
specimens was taken in swampy riverine forest, one in open forest under-
growth, and one in dense undergrowth of mature forest.

1978

The Lathrop Ornithological
Central African Expedition

5

Family Columbidae

Turtur tympanistria tympanistria (Temminck and Knip). — One male,
testes enlarged, 5 June, weight 67 g.

Turtur brehmeri brehmeri (Hartlaub). — The Blue-headed Dove must be
fairly common in the dense forest around the Ouossi River as 4 examples were
obtained on four different days, 1,2, 10 and 16 June. Two of them had enlarged
gonads, the other two had small, inactive ones; 1 male weighed 120 g; the 3
females 105, 107 and 130 g respectively.

Family Cuculidae

Clamator levaillantii Swainson. — This crested cuckoo is not a bird of the
true forest, but does come into the margins of such areas. One adult male was
captured in a mist net in the undergrowth of mature forest 8 June; testes
somewhat enlarged; weight 110 g.

*Pachycoccyx audeberti (Schlegel). — Because of the still limited number
of records of this scarce cuckoo, it may be noted that Andrew Williams saw,
but was unable to collect, one just outside the forest.

Cuculus solitarius solitarius Stephens. — One adult male, testes not en-
larged, was taken in open forest, bordering grassland, 7 June; weight 83 g.

*CucuIus clamosus gabonensis Lafresnaye. — One adult female, ovary
enlarged, was taken in open forest 8 June; weight 78 g. This specimen, together
with one of C. solitarius, collected the day before, raise again the old doubts as
to the nature of the relationship of the two species. Furthermore, Williams
wrote that he also saw, but did not obtain, one pure black clamosus. Were it
not for the fact that in their vocalisms, the coloration of their egg shells, and
their different choice of hosts, clamosus and solitarius, at least in southern and
eastern Africa, are quite distinct, one might be tempted to regard them as
variables within a single broadly phenotypic species. This is, however, negated
by these considerations and the only possible conclusion is that the two species
may have had a common origin in the forested regions of central Africa where
the persistent, present population still reveals the phenotypic spectrum of the
ancestral stock (including such variables as jacksoni, gabonensis and mabirae).
It is known to occur, not far to the south, in the forests of Zaire. The all black
individual seen by Williams must have beenC. clamosus clamosus of southern
Africa, a race known to migrate north after the southern breeding season to the
Congo forests and even as far north as southern Ethiopia. It too, is a first
record for the Central African Republic.

*Cercococcyx mechowi Cabanis. — The Dusky Long-tailed Cuckoo is gen-
erally thought of as an uncommon, or, at least, elusive bird, “ . . . more easily
heard than seen” (Mackworth-Praed and Grant, 1970:369), of the central Afri-
can forests, but it must be anything but scarce in the Ouossi River area, as five
examples, all females, were obtained there, 1, 10, 12 and 17 June. One of these
was an immature bird, the others were adult; some were shot from as high as 40

6

Contributions in Science

No. 287

feet up in trees while others were taken in mist nets set in the undergrowth. The
weights of the adults were 54, 55, 59 and 61 g; the young bird 61 g. One bird had
the ovary enlarged; the others had small ovaries.

Family Caprimulgidae

Scotornis climacurus sclateri Bates. — One adult male, testes enlarged, 5
June, weight 44 g, taken in open country near the forest edge.

Family Trogonidae

Apaloderma narina brachyurum Chapin. — One adult male, testes slightly
enlarged, weight 63 g, 10 June, was collected in mature riverine forest with
open undergrowth.

Family Alcedinidae

Alee do quadribrachys guentheri Sharpe. — Four specimens, 10 to 18
June, were taken in mist nets over a small stream in riverine forest; all had
small, inactive gonads; 2 males weighed 33 and 36 g, 2 females 32 g each.

*Alcedo leucogaster leopoldi (Dubois). — Between 6 and 16 June, 4 males
and 3 females were taken in mist nets set in swampy riverine forest. The birds
had gonads ranging from not enlarged to slightly so; weights were 12, 12, 12 and
14.5 g for the males; 14, 14 and 16 for the females. The present specimens agree
in coloration and in size with a series from western Uganda, but show less
diversity in the purplish or greenish-blue bars on the coronal feathers. In the
birds from two forests in Uganda, the Bwamba in extreme western Uganda,
and the Malabigambo near the western shore of Lake Victoria, it was found
(Friedmann 1969:4) that there were almost two color phases in this character
and that there were no intermediates. This is not the case with the Ouossi
River birds, some of which actually have both purplish and greenish-blue bars
on different feathers.

Myioceyx lecontei (Cassin). — Between 6 and 15 June, 6 specimens of the
Dwarf Kingfisher were netted in dense riverine forest undergrowth. None
were in breeding condition; weights 10 to 10.5 g (4 males), 10 and 12 g (2
females). Comparison of these specimens with a long series in the Los Angeles
County Museum from western Uganda, including a number from the Budongo
Forest (type locality of van Someren’s proposed race M. /. ugandae ) show no
differences in dimensions or in coloration. The small blue coronal spots that
were the basis of ugandae are by no means constant in Uganda birds, and are
present in 2 of our 5 adults from Ouossi River. I conclude that this species has
no recognizable races.

Ispidina picta picta (Boddaert). — The Pigmy Kingfisher is represented by
2 males and 1 female taken 16 and 19 June in mist nets set in swampy riverine
forest; all in non-breeding state, or, at most, with slight gonadal enlargement;
weights 12 and 12.5 g (males); 11 g (female).

1978

The Lathrop Ornithological
Central African Expedition

7

*Halcyon badia Verreaux. — One female, ovary not enlarged, was netted
in the forest undergrowth 18 June. Comparison with a dozen west Uganda
specimens in the Los Angeles County Museum, including a series of topotypes
of budongoensis from the Budongo Forest, shows no reason for recognizing that
subspecies. It is sufficient to note a general clinal increase in size toward the
eastern parts of the range of the species. Our specimen weighed 47 g. This
specimen was caught in mist nets which suggest that Mackworth-Praed and
Grant (1970:431) were in error when they wrote that this species usually
“. . . keeps twenty or thirty feet from the ground.”

*Halcyon malimbica malimbica (Shaw). — Four specimens, 2 of each sex
were taken (3 in mist nets) in the riverine forest undergrowth, 1 to 15 June, all
with slightly enlarged gonads; weights 83 and 90 g (males); 83 (both females).

Family Bucerotidae

*Tockus hartlaubi grand (Hartert). — A breeding male, one of two birds
seen about 60 feet up in a tree, catching insects on the wing, was taken 3 June in
lush forest; weight 135 g. This specimen agrees in coloration with another male
in the Los Angeles County Museum from Bwamba Forest, western Uganda, but
has a much longer bill (culmen to base 74 mm; 63 in the Bwamba bird). There is
some difference in the wing and tail lengths of the two, but the Uganda bird is in
very abraded plumage, especially in its remiges and rectrices, which naturally
affects its dimensions.

Family Capitonidae

*Pogoniulus scolopaceus flavisquamatus (Verreaux). — One female,
ovary not enlarged, weight 13 g, was netted in dense undergrowth in degraded
forest, 5 June.

Pogoniulus chrysoconus chrysoconus (Temminck). — Three male exam-
ples of the Yellow-fronted Tinker-bird were collected, 17 and 19 June, in mist
nets set at a fruiting tree at the forest edge; weights 10, 1 1 and 11.5 g; one of
them had large testes, the others small ones. They agree in size and coloration
with other specimens in the Los Angeles County Museum from the Ivory Coast,
Uganda and western Kenya.

"Pogoniulus bilineatus leucolaema (Verreaux). — Four males, 2 with
large, 2 with small gonads, were collected 8 to 18 June in the forest edge;
weights 10.5 to 12 g (average 11.1). These specimens are slightly paler below
than a long series of P. b. mfumbiri, less yellowish below than P. b. sharpei
from the Ivory Coast.

* Pogoniulus sub sulphur eus flavimentum (Verreaux). — The Yellow-
throated Tinker-bird was common in the Ouossi River area, where 3 females in
non-breeding state, and 2 males with enlarged testes, were collected 15 to 17
June; weights 9 and 10.5 g (males); 10 g (females). All were collected while
feeding in fruiting forest trees. These specimens agree in size and coloration
with a long series in the Los Angeles County Museum from western Uganda.

8

Contributions in Science

No. 287

*Pogoniulus atroflavus (Sparrman). — The Red-rumped Tinker-bird was
found to be very common in the Ouossi River area, and some 13 specimens
were collected 12 to 19 June; 6 males, 7 females, gonads enlarged in 4 individu-
als, small in the others; weights 17 to 21.5 g (average 18.1) in males, 14 to 20 g
(average 17.5) in females. Three of the birds were caught in mist nets, but the
majority (10) were shot as they were feeding well up in fruiting trees.

*Trachyphonus purpuratus purpuratus Verreaux. — The Yellow-billed
Barbet was met with twice, 9 and 16 June, in the riverine forest; 1 male with
slight testicular enlargement and 1 female with a small ovary; weights 87 g
(male), 89 g (female). These examples are placed with the nominate race of the
species, but it is not clear that T. p. elgonensis is constantly different. That
eastern subspecies is said to have the whitish edges of the feathers of the throat
and foreneck less well marked, but a very long series in the Los Angeles
County Museum from the forests of western Uganda suggests that this charac-
ter is one that appears as the feathers become abraded. Some of the Uganda
birds have these edges as well developed as in our Ouossi birds, while others
have no sign of them. Unfortunately, we have insufficient west African mate-
rial for comparison, but it would seem that only specimens in fresh plumage of
the two can be expected to demonstrate the racial difference. The distribu-
tional maps in Mackworth-Praed and Grant (1970:553) suggest that they had no
records from the area we are concerned with in this report, but their range for
T. p. purpuratus comes nearer to it than that of T. p. elgonensis. White
(1965:271) writes that the nominate race intergrades in the upper Congo with
elgonensis, and it would seem that this is true as well in the Central African
Republic.

Family Indicatoridae

*Melignomon zenkeri Reichenow. — One specimen of this rarely col-
lected species was caught in a mist net 1 m above the ground in dense forest
undergrowth by a small stream along the Ouossi River, 2 June; a female in
breeding condition, the largest ovarian follicle 2.5 mm; bill dark horn brown,
the lower base yellowish; iris dark brown; feet pale olivaceous yellow; eye skin
dark gray; weight 24 g. The specimen agrees in coloration with another breed-
ing female from Bwamba (Los Angeles County Museum #66898), extreme
western Uganda (July 9), but is smaller, this being especially noticeable in the
bill and tail. The Ouossi bird measures: wing 76, tail 47.6, culmen from the base
10.1 mm; the Bwamba one: wing 77.6, tail 53.7, culmen from base 11.8 mm,
weight 25 g.

The stomach of the present specimen contained a sizeable quantity of
finely ground or comminuted grayish waxy material mixed with tiny black bits
of insect fragments, very similar to that reported in detail from the earlier
Bwamba specimen (Friedmann 1968:281-282), which was determined to be
scale insect wax, not the paler, more yellowish beeswax usually present in the
stomachs of honey-guides of the genus Indicator.

1978

The Lathrop Ornithological
Central African Expedition

9

The fact that the genus Melignomon is intermediate in its characters be-
tween Indicator and Prodotiscus, but somewhat more like the latter, makes it
of interest to note that while taking the present specimen out of the mist net,
Andrew Williams found it to have a very “hard” body like that of an Indicator,
not like the softer, “delicate” body of a Prodotiscus.

* Indicator maculatus stictithorax Reichenow. — The Spotted Honey-
guide was found to be very common in the forests along the Ouossi River, and
16 males, 17 females, and 1 unsexed specimen were collected, 31 May to 19
June. While most of the birds had no, or only little, gonadal enlargement, 2
taken on 31 May were marked as breeding and 1 other, collected 3 June, had an
enlarged ovary. Most of the specimens were caught in mist nets placed near
wild bees’ nests, baited with bits of beecomb. The weights of the birds varied
from 43 to 51 g (average 50.6) in males; 40 to 50.5 g (average 40.6) in females.
Almost all the birds had beeswax in their stomachs, many also had insect
fragments, and one had 2 small seeds as well as insect parts. The condition of
the plumage varies from extremely worn to fresh, but no correlation between
gonadal state and feather wear could be discerned.

The above series presented an opportunity to compare central African
with western Ugandan (Bwamba) birds. The more eastern population averages
slightly larger, but the limits of size variation are about the same in our
Bwamba and Ouossi River birds (no topotypical stictithorax, described from
Cameroon, have been available for direct comparison). Thus, males have wing
lengths of from 95.5 to 106.5 mm (average 101 .5) in 12 Bwamba specimens, 95.0
to 104 mm average (99.9) in 17 Ouossi River birds; females have wing lengths
of 95 to 98.8 mm (average 96.4) in 1 1 Bwamba birds, 92.1 to 100.3 mm (average
90.4) in 16 Ouossi River examples.

A general clinal increase in size occurs in the species as a whole from west
to east. Mackworth-Praed and Grant (1970:558) write that nominate maculatus
(Gambia to Nigeria) have wing lengths of 97 to 103 mm, stictithorax 100 to 107
mm. I have found no specimen with wings as long as 1 10 mm, the maximum for
stictithorax given by Malbrandt and Maclatchy (1949:275).

According to Chapin (1939:548) typical maculatus differs from stictithorax
in being a little darker on the crown, the cheeks and malar region unstreaked,
dark olive, and the abdomen less yellowish. Judging from our large series of
stictithorax it would seem that the abdominal coloration is too variable to be of
much diagnostic value, but all of our 59 birds have streaks on the cheeks and
malar region.

Indicator indicator (Sparrman). — The Greater Honey-guide is repre-
sented by 4 adult males, 1 adult female, 3 immature birds of both sexes, taken 1
to 15 June, all with little or no gonadal enlargement, weights 45 to 50 g (average
48) in the adult males, 52 g in the adult female; 45 and 46 g in 2 immature
females, 54 g in an immature male. The specimens were taken at the edge of the
forest, some of them in mist nets near wild bees’ nests; they were in remarka-
bly fresh, unfaded plumage.

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Clancey (1970:378) described a race of this honey-guide from southern
Mozambique, I. indicator inquisitor, said to be characterized by smaller size;
wing length of males 105 to 110.5 mm, as against 113 to 120 in males of the
nominate subspecies. While the validity of a southeast African race may seem
tangential to a study of a series from the Central African Republic, it does
affect the present use of a binomial for our specimens. Our males from Ouossi
River have wing lengths of from 104 to 107.8 mm (average 106.4), and our
comparative material from Kenya have wings 1 03 to 1 09.8 mm ; in other words all
are within the size range given for the race inquisitor; one male from the Ivory
Coast has a measurement of only 102.5 mm. The wide geographic spread of
small /. indicator makes one ask whether typical indicator may be a large race
confined to the Cape Province, an area from which no material has been
available for comparison. However, Clancey stated that coastal Kenya birds are
equally large and must be considered /. indicator. Our coastal Kenya adult
males (Lamu, Tana River and Sokoke) have wing lengths of from 104.5 to 107.0
mm. The mensural data given by Irwin and Benson (1966:16) for birds from
various parts of Zambia are similarly difficult to account for in terms of
geographic subspeciation, although in a later paper Clancey (1972:181-182)
extended the range of inquisitor to include . . the valley of Luangwa, Zambia,

and that of the lower Shire R., Malawi, and, perhaps, on the coast of Tanzania
and the off-shore islands of Mafia, Zanzibar and Pemba.” However, it seems
unlikely that this supposed race could also extend to the Central African
Republic and even to the Ivory Coast, and until further clarification, it seems
better to consider the species monotypic.

Indicator minor riggenbachi Zedlitz. — The Lesser Honey-guide was
found to be very common along the Ouossi River in open areas at the edges of
the forest; 12 males and 3 females were collected 3 to 18 June; all with little or
no gonadal enlargement; weights 27.5 to 34 g (average 30.4) in the males, 27 to
30 g (average 28.3) in the females. Many of the specimens were caught in mist
nets near wild bees’ nests, and most (1 1) of them had beeswax as well as insect
fragments in their stomachs. This series agrees very closely with many others
from extreme western Uganda, and does not show any trend toward the ventral
pallor of I. minor alexanderi, the race that occurs to the northwest of their area,
and still less to the still paler I. minor senegalensis with which White (1965:275)
has united alexanderi.

* Indicator exilis exilis (Cassin). — The Least Honey-guide must be a
very common bird in the forest along the Ouossi River, as in 17 days, 2 to 19
June, the expedition obtained a series of 29 specimens (17 males, 12 adult, 5
immature; and 10 females, 6 adult, 4 immature; and 2 birds of unrecorded sex).
As might be expected from the geographic location of the area, the birds are
somewhat intermediate between nominate exilis and the slightly larger, more
eastern, pachyrhynchus (Bahr-el-Ghazal province of the Sudan to eastern
Zaire, Uganda and western Kenya), but seem better placed with the former
subspecies. Thus, the wing lengths of our 12 adult males vary from 68 to 80.5
mm (average 75.9); 6 adult females 66.2 to 71.2 mm (average 68). Chapin

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(1962:41-42) noted wing lengths of 72 to 79.5 mm for male exilis, 65 to 70 for
females; while for pachyrhynchus he gave wing lengths of 75.5 to 84 mm for
males, 68 to 76 mm for females. In the Itombwe area of eastern Zaire,
Prigogine (1971:101-102) found the two races to be altitudinally distinct. I. e.
exilis below 1200 m, and I. e. pachyrhynchus above 1800 m, a conclusion
difficult to accept. In this connection I have reviewed and measured all the
specimens in the Los Angeles County Museum collections from the Impenetr-
able Forest, southwest Uganda, and find them all to be pachyrhynchus; they
were collected at altitudes of from 4000 to 7500 feet. In the report on the birds
of the Impenetrable Forest (Keith, Twomey, Friedmann, and Williams
1969:14) only /. exilis exilis is listed and only from 4000 feet. Some of our
specimen records from higher elevations were collected too late for inclusion in
that paper, and at that time pachyrhynchus was considered not different
enough from exilis to recognize by name. The two races are based, it is true, on
only average differences; Mackworth-Praed and Grant (1970:566) also consider
them as one subspecies. However, in view of Chapin’s long experience with
these birds in the field and museum, I am inclined to follow him in keeping the
two apart.

The 10 immature examples of our present series are noticeably duskier
below than the adults, and lack the black malar stripe and the white loreal
streak of the older birds. Identical differences between young and adult birds
are also present in a still larger series of specimens from Bwamba and other
forests of extreme western Uganda. The young birds average slightly smaller
than the adults; their wing length averages 72.5 mm in the males, 67 mm in the
females. Of the 12 adult males in our Ouossi River series, none had testes more
than slightly enlarged; of the 6 adult females, all but 1 had enlarged ovaries, an
anomalous situation.

Many of our specimens were collected in mist nets near wild bees’ nests at
the edge of the forest, but a few were shot while feeding on fruits well up in
small trees. Most of them had beeswax and insect fragments in their stomachs,
but 2 of them, both immature, had small fruits as well. This is the first time this
Honey-guide (or any of its congeners) has been found to eat small fruits, an
unexpected diet for a Honey-guide. In my earlier (1955:227) survey of the
feeding habits of this species, I noted that a seed of unknown kind had been
found in the gizzard of one specimen, but otherwise the stomach contents were
invariably insects and beeswax (and probably honey). The 12 adult males
weighed from 16 to 20.5 g (average 18.2), the adult females 12 to 17.5 g (average
15.9); immature males 16 to 18 g (average 17.5); immature female 17 g.

The use of mist nets placed near bees’ nests as a method of obtaining
specimens of this Honey-guide was well documented by Archer and Glen
(1969:1-2) in their work in the Malabigambo Forest, Uganda. They concluded
that their success was due to the fact that the birds made periodic visits to all
the wild bees’ nests known to them. Despite there being a total of over 2500
yards of mist nets used in the month’s work in that forest, not a single Honey-
guide was netted except in the immediate vicinity of the bees’ nests, although

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in the Namalala Forest they did catch one in a net "set at random" (but
possibly not very far from a bees' nest unknown to the collectors).

The receipt of this fine series, added to the great number of specimens
from western Uganda, now in the Los Angeles collections, has caused me to
restudy the variation of I. exilis. This leaves me with serious doubt as to the
identification of one example (LACM #66903) from Ntandi, Bwamba Forest,
Uganda, previously published (Friedmann and Williams 1968: 1 8 ; 197 1:33) as I.
pumilio. It now seems that minimal specimens of exilis are just as small as
pumilio, and indeed, can hardly be distinguished from the latter, without care-
ful study. Thus, the Ntandi bird has a bill slightly larger than in pumilio, more
like exilis, to which species it is now identified even though it is actually
exceeded by some examples of pumilio in its wing length, tail length and body
weight. The minute bill of the latter species seems to be its most reliable diag-
nostic character; to be appreciated, the bills of exilis and pumilio should be
examined with a hand lens, preferably from the underside. Although the
specimen from Kakamega Forest, western Kenya, that Williams and I previ-
ously (1965:21-22) decided was I. pumilio, is not available for restudy, I began
to wonder if it too might be a minimal I. exilis. However, Prigogine writes me
(25 July 1977) that he recently examined it and one other Kakamega specimen
in the Nairobi museum, and found them to be I. pumilio, of an undescribed
subspecies, which, I hope, he will elucidate in his next paper. Zimmerman’s
report of pumilio from the Kakamega Forest (1972:295) was based solely on a
sight record.

While collecting the present series, Williams had in mind the possibility of
some of them being/, willcocksi, as I also had when studying them, but all are /.
exilis.

*Melichneutes robustus (Bates). — The Lyre-tailed Honey-guide is still so
scarce a bird in museum collections (not, apparently, in nature) that additional
specimens are examined with much interest. Two adults, one of each sex, were
caught in mist nets set in open forest near a wild bees’ nest, and baited with
beecomb, 6 and 9 June. The male showed some gonadal enlargement, testes 4
x 3 mm, the female had a small, inactive ovary. The female was in much worn
and faded plumage; the male was not in fresh, but in less abraded, feathering,
and was darker above; weight 61.5 g in the male, 49.5 in the female; stomach
contents of male "entirely beeswax — weighed 3 g"; of female "insect frag-
ments"; soft parts were recorded as follows: bill dark horn brown, lower base
paler; iris orange-brown (male) to dark brown (female); feet dark brownish gray
to blackish gray; bare eye skin pinkish brown (male), olive gray (female); wing
length 95.8 mm (male), 94 mm (female).

Inasmuch as so little is still known of this Honey-guide it may be well to
call attention to a specimen taken more than 12 years ago, in a snare baited with
beecomb, in the Mongomo region, Rio Muni (now Spanish Continental
Guinea), a record published anonymously (probably by J. Sabater Pi) in a
little-consulted popular journal of the Barcelona Zoological Park (Anon.
1963:43). This record has been overlooked by all the ornithological journals,
and is the first, and so far, the only, record from that country. The specimen is

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now in the American Museum of Natural History (AMNH #802455). While this
record did not increase the known range oiMelichneutes, which was previously
reported from Gabon to the south, and from Cameroon to the north, the almost
simultaneous report by Beatty (1963:100-101) suddenly extended its range
westward by about 1000 miles to Mt. Nimba, at the point where Liberia, Ivory
Coast and Guinea come together. The previous westernmost record was in
southern Nigeria, but it is probable that the range is not that discontinuous; the
species may be expected to “turn up” in the upper Guinean forests of the
Ivory Coast, Ghana and Dahomey.

Family Picidae

*Campethera nivosa herberti (Alexander). — The Buff-spotted Wood-
pecker was found to be common along the Ouossi River and 4 males and 3
females were collected 1 to 18 June, all with little or no gonadal enlargement;
weights males 34 to 38 g (average 37); females 34.5 to 38 g (average 36.1).

Campethera caroli (Malherbe). — One male, 4 females, all with sjnall
gonads, taken 6 to 17 June; weights: male 60 g; females 53 to 68 g (average
58.6). These specimens agree with a long series from western Uganda in the
Los Angeles collections, and would have to be considered budongoensis if that
race were upheld, as it is by Mackworth-Praed and Grant (1970:571), but the
characters of that subspecies do not appear to be constant enough to warrant
its recognition.

Dendropicos xantholophus Hargitt. — This woodpecker apparently is less
common in the area than either of the Campethera species; only a single
example was obtained 5 June, a male with small testes; weight 58 g. It was shot
in the tree tops, whereas the Campethera were taken largely in the mist nets,
which may account for their apparent greater numbers and relative abundance.

Family Eurylaemidae

*Smithornis capensis camarunensis Sharpe. — Three specimens, 1 saved
as a study skin and 2 as skeletons, were collected in mist nets in the forest
undergrowth 2 to 13 June; the male preserved as a study skin and a female
saved as a skeleton had enlarged gonads; a male (skeleton) showed only slight
testicular swelling; weights 26 g in each male, 24 g in the female. The subspecific
identification of the study skin is based on the conclusions of Mackworth-Praed
and Grant (1970:609), but it agrees very well with a series of meinertzhageni
from western Uganda, which Mackworth-Praed and Grant consider the same as
medianus. More extensive series with more complete geographic coverage are
needed to settle the matter of races of this bird.

Family Pittidae

*Pitta angolensis longipennis Reichenow. — One male, testes not en-
larged, was taken in a mist net 3 feet up in dense undergrowth of mature forest,

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5 June, wing 121.5 mm, weight 84 g. The fact that this bird, and also its other
African congener, P. reichenowi, are captured so seldom, even in mist nets,
suggests that they are really scarce and not merely difficult to see.

Family Alaudidae

Mirafra rufocinnamomea tigrina Oustalet. — The Flappet Lark is a bird
of the open grasslands, not of the forest; 2 males, 1 with enlarged, the other
with small testes, were taken just outside the forest edge 6 and 16 June; weights
27 and 28 g. This race of the Flappet Lark has been collected but seldom; no
records from the Central African Republic for the species are indicated in Hall
and Moreau (1970:map 6). Meyer de Schauensee (1949:8) listed a specimen of
M. r. zombae from Oka, which may be referable to tigrina. White (1961:20)
places zombae in the synonymy of fischeri, but it seems likely that the name
tigrina could be applied to a specimen from the Ubangi-Shari region.

Galerida modesta bucolica (Hartlaub). — The Sun Lark was found in the
open grasslands at the edge of the forest; 2 males and 1 female, all with small
gonads, were taken 6 and 12 June; weights 21.5 and 22 g in the males, 18 g in the
female; stomach contents, insect fragments and grass seeds.

Family Pycnonotidae

Pycnonotus virens virens (Cassin). — The Little Greenbul was common in
the Ouossi River area; 5 specimens were netted in the forest undergrowth 31
May to 9 June, 3 males with enlarged testes, and 2 females, one with enlarged,
the other with slightly englarged ovary; weights males 23.5, 24 and 24 g;
females 21 g.

Pycnonotus curvirostris curvirostris (Cassin). — Fairly common; 3 males
and 1 female were netted in the riverine forest undergrowth 2 to 17 June;
gonads enlarged in the 3 males, slightly so in the female; weight males 24, 25,
26 g; female 26 g. Only one record from the Central African Republic is indi-
cated in Hall and Moreau (1970:map 68); none in Malbrandt and Maclatchy
(1949).

Pycnonotus latirostris latirostris (Strickland). — Two females, 1 with
small, 1 with enlarged ovary, were netted in the forest undergrowth 31 May and
10 June; weights 22 and 30 g. Because these birds are fairly small, wing length
72 and 76 mm, they are placed with the nominate race of this Greenbul, in
accordance with Chapin’s (1953:113) arrangement, although they are not
smaller than some of a long series of eugenia from Uganda. Chapin admitted
that it was “difficult to determine the limits’’ between the two. In the Central
African Republic this species must be restricted to the forested areas in the
south along its border with the Republic of the Congo.

"Pycnonotus gracilirostris chagwensis (van Someren). — One bird of each
sex, both with gonads enlarged, shot in a fruiting tree at the forest edge 8 and 15
June; weight 32 g (male), 40 g (female). The map in Mackworth-Praed and
Grant (1970:82) seems to anticipate its presence there, but as stated earlier, this
is not a definite statement.

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*Baeopogon indicator indicator (van Someren). — One male, testes
slightly enlarged, June 19; weight 46 g.

Phyllastrephus scandens orientalis (Hartlaub). — Six specimens, 8 and 9
June, gonads enlarged in 1 male, not enlarged in 2 others and in 3 females;
mostly netted in open forest undergrowth; weights males 48 and 50.5 g, females
38, 41, 45 g; all the specimens in worn plumage.

Phyllastrephus albigularis albigularis (Sharpe). — Common; 6 specimens
were caught in mist nets in the forest undergrowth 31 May to 10 June; 3 with
large, 3 with small gonads; weight males 26 and 28.5 g, females 20 to 24 g
(average 22.2). These birds agree in size and coloration with a long series from
western Uganda. Ouossi River provides a specimen record locality that fills a
large gap in Hall and Moreau’s map (1970:map 90).

Bleda syndactyla woosnami Ogilvie- Grant. — Five adults, taken in
riverine forest undergrowth 31 May to 7 June; gonads varying from small to
large; weights males 48, 49 g; females 41, 48 g. These specimens agree with a
long series from western Uganda and do not show any intergradation with B.
syndactyla multicolor, although the distribution maps in Mackworth-Praed and
Grant (1970:57-58) suggest that the latter might be the race to be expected in
the southern part of the Central African Republic. Hall and Moreau (1970:map
80) show only one record of this bird in the Central African Republic, where its
range must be limited to the southern forests near the border of the Republic of
the Congo.

Bleda eximia ugandae van Someren. — The Green-tailed Bristle-bill is
represented by 5 adult males, testes large in 4, slightly enlarged in 1 , and 1 adult
female, ovary slightly enlarged; 31 May to 7 June; weights 42 to 48 g in males
(average 44); 36 g in the female.

Nicator chloris (Valenciennes). — Although many recent authors place
Nicator in the shrike family, I prefer to follow Chapin’s arguments (1953:183-
185) and keep it among the bulbuls, somewhat aberrant as it may be in that
family. It must be common in the Ouossi River forests, where 7 adults were
obtained, largely in mist nets in the undergrowth in both open and dense forest,
2 to 19 June; 3 males with enlarged testes, 4 females with small or slightly
enlarged ovaries; weight males 50, 54, 58 g; females 40, 41, 43, 43g.

Criniger calurus emini Chapin. — Fairly common; 5 specimens, 3 males
and 2 females were taken in mist nets in the riverine forest undergrowth, 3 to 16
June; gonads not enlarged in 3, enlarged in 2 of the birds; weights males 30.5 to
34 g (average 32.1); females 25 and 30 g.

Family Laniidae

Malaconotus cruentus adolfifriederici Reichenow. — One male, testes
slightly enlarged, was taken in dense forest undergrowth 31 May; weight 79.5
g. The races of this shrike are ill-defined and not too certainly identifiable.
However, the small size of the present specimen, wing 107 mm, agrees with
one from the Semliki River (Chapin 1954:40), and it is placed with adolfi-
friederici. Hall and Moreau (1970:map 112) show no records for M. cruentus

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from the Central African Republic east of Bangui, and Mackworth-Praed and
Grant (1973:460) seem to restrict adolfifriederici to the eastern parts of Zaire
and western Uganda, so the present specimen extends the known range of that
race a long distance to the northwest.

Family Muscicapidae
Subfamily Turdinae

*Erythropygia leucosticta collsi Alexander. — Three adult males, 1 breed-
ing, 1 with enlarged and 1 with small testes; 1 female with an enlarged ovary,
were taken in mist nets in heavy forest undergrowth 6 to 12 June; weights males
26, 27, 28 g, female 27 g; stomach contents beetles and a small snail.

Erithacus erythrothorax mabirae Jackson. — A common bird in the
Ouossi River forest, but one that is seldom seen except when caught in mist
nets; 5 males and 2 females were so captured in dense forest undergrowth 31
May to 4 June, gonads large in 1 male, small or only slightly enlarged in the
others; weights males 16.5 to 19 g (average 17.5), females 16.5, 17 g. Judging
from the absence of records in Hall and Moreau (1970: map 145) the above are
the first specimens collected in the Central African Republic other than in the
extreme southwest, near the Cameroon border.

*Cossypha natalensis intensa Mearns. — Two males, 1 female, all with
gonads much enlarged, taken in forest undergrowth 8 to 10 June; weights males
37, 39 g, female 38 g. These extend the known range of this Robin-chat
northward to the southern part of the Central African Republic.

*Alethe diademata woosnami Grant. — The Fire-crest Alethe was found
to be common in the Ouossi River forests and its range in the Central African
Republic is probably restricted to the forests near the Zaire border. Between 31
May and 9 June 8 specimens were taken, 4 males with large testes, 1 female
with an oviduct egg, 2 with slight ovarian enlargement, and 1 subadult female
molting into adult plumage; weights males 32 to 34 g (average 32.6), female
adults 32 to 39 g (average 34.7), subadult female 28 g.

*Alethe poliocephala carruthersi Grant. — Four adult Brown-chested
Alethes, two of each sex, the males with large testes, the females varying from
small to slightly enlarged ovaries, were netted in the dense forest undergrowth
31 May to 15 June; weights males 30, 32.5 g; females 26, 31 g respectively.

Stizorhina fraseri vulpina Reichenow. — Fairly common in the Ouossi
River forests, 1 male and 4 females were taken there 1 to 11 June; gonads
enlarged in most of the birds, one female marked “breeding”; weight of male
34 g, females 32, 33.5, 36 and 39.5 g; all had been feeding on small beetles.

Neocossyphus poensis praepectoralis Jackson. — Two males, 1 female,
with gonads enlarged in 1, not in the others, were taken in riverine forest
undergrowth 31 May to 10 June, and extend the known range of this bird a
considerable distance north. In the Central African Republic the species was
reported earlier only from the extreme southwest corner, close to the Came-
roon border. The males weighed 52 and 53 g, the female 50.5 g.

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Subfamily Timaliinae

*Trichastoma albipectus albipectus (Reichenow). — Common in the
forest undergrowth; 4 males and 3 females were taken 1 to 19 June, gonads
varying from small to large; weights males 31 to 38 g (average 35.2), females 30
to 36 g (average 32.3); stomach contents beetles and one lepidopteran larva.
These specimens extend the known range of the species to the northwest of the
records plotted by Hall and Moreau (1970:map 165).

*Trichastoma rufipenne rufipenne Sharpe. — Apparently less numerous
than T. fulvescens; 3 males, all with enlarged testes, taken 2 to 5 June, weights
26 to 30 g (average 28), agree with a long series from western Uganda.

Trichastoma fulvescens ugandae (van Someren). — A common species in
the forest, 4 adults of each sex were collected 1 to 17 June; gonads varying from
not enlarged to large; weights 31 to 38 g (average 34.7) in males; 28 to 30.5 g
(average 29) in females.

*Ptyrticus turdinus turdinus Hartlaub. - — Two examples of the Thrush-
babbler were taken in the dense undergrowth of the forest 1 1 and 12 June, one
of each sex, both with small gonads; weight male 72.5 g, female 59 g. The male
is considerably larger than the female, wing 107.4 as against 95.4 mm; culmen
from base 22.9 as against 22.1 mm; the tail of the female too damaged for
meaningful comparison. The present specimens help to fill a great geographic
gap in the range, as indicated by actual specimen records, in Hall and Moreau
(1970:map 168).

Phyllanthus atripennis bohndorffi (Sharpe). — The Capuchin Babbler is a
common bird in the forests of the Ouossi River area. Six adults, 3 of each sex
were taken between 5 and 16 June (4 netted, 2 shot); all with small or only
slightly enlarged gonads; weights males 80, 84, 90 g; females 86, 86 and 94 g
respectively. These specimens are all typical bohndorffi and show no approach
to haynesi of northern Cameroon although 1 male has two dark brown feathers
among the gray ones on one side of the crown; they agree closely with a series
from western Uganda. The Ouossi region adds an intermediate locality of
record in the distribution shown for the species in Hall and Moreau (1970:map
175).

Subfamily Sylviinae

Cisticola natalensis strangei (Fraser). — Two males, testes slightly en-
larged, were taken at the forest edge 15 and 20 June; weights 23 and 26 g.

Camaroptera brevicaudata tincta (Cassin). — Three males, all with large
gonads, were netted in the forest undergrowth 4 to 12 June; weights 1 1, 12 and
12 g.

* Camaroptera superciliaris flavigularis Reichenow. — One male, testes
large, was taken in dense growth at the forest edge 4 June; weight 10 g.

* Camaroptera chloronota toroensis (Jackson). — Three males, testes
slightly or not enlarged, netted in forest undergrowth 1 and 2 June, weights
10.5, 1 1 and 12 g, constitute a very considerable northwestern extension of the

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known range of the species. They agree well with west Ugandan examples and
are, accordingly, placed with that race.

*Macrosphenus flavicans hypochondriacus (Reichenow). — One male
with small testes, netted in forest undergrowth 10 June, weight 14 g, extends
the previously known range of the race hypochondriacus northwestward about
200 miles.

Hyliota flavigaster flavigaster Swainson. — Two adults, one of each sex,
male with enlarged testes, female with small ovary, were collected at the edge
of the forest 12 and 13 June, foraging in the tree tops with a party of Parus
leucomelas; weight male 13 g, female 12.5 g. Judging by the distribution shown
in Hall and Moreau (1970: map 259) these specimens fill a considerable gap in
the recorded range of the species.

Hylia prasina prasina (Cassin). — Apparently common in the Ouossi
River area, 6 specimens, 3 of each sex, gonads varying from small to large,
were netted in the forest undergrowth 1 to 10 June; weights males 14, 15, 15 g,
females 12 g in each case.

Subfamily Muscicapinae

Fraseria cinerascens cinerascens Hartlaub. — This Flycatcher must be
common in the Ouossi River forests, as 4 adults of each sex and 1 immature
male were taken, chiefly in mist nets in the forest undergrowth, 31 May to 18
June; gonads small in most birds, large in 1 male taken 31 May; weights males
21.5 to 24.5 g (average 23), females 18 to 22 g (average 19.8).

Myioparus plumbeus plumbeus (Hartlaub). — A single male, testes not
enlarged, one of a small flock seen at the edge of the forest, was collected 18
June. It is not fully adult, with a brownish wash on the breast and sides of the
abdomen; weight 13 g. Hall and Moreau (1970:map 249) show no records for
this Flycatcher from the southern part of the Central African Republic.

Myioparus griseigularis griseigularis (Jackson). — Three males, 2 with
enlarged testes, 1 with small ones, 2 to 12 June; weights 14 g in each case;
plumage somewhat abraded in all. These specimens agree closely with a series
from western Uganda. The Ouossi River records add a locality in the middle of
a considerable gap in the map in Hall and Moreau (1970:map 249).

*Platysteira castanea castanea (Fraser). — Two males and 1 female, all
with enlarged gonads, were netted in dense forest undergrowth 4 and 9 June;
weights males 13 and 14 g, female 16 g.

*Trochocercus nigromitratus (Reichenow). — One female, with ovary not
enlarged, was taken in open undergrowth in riverine forest 6 June; weight 9 g.
The distribution of this species in the Central African Republic must be limited
to the forested areas of the south, along the border of the Republic of the
Congo.

Terpsiphone rufiventer ignea (Reichenow). — Apparently common in the
Ouossi River area, but previously reported only from the western part of the
Central African Republic; 4 males, all with large testes, 2 females, with little or

1978

The Lathrop Ornithological
Central African Expedition

19

no ovarian swelling, 7 to 18 June; weights males 14 to 15.5 g (average 14.9),
females 15.5, 16 g. These specimens extend the known range of ignea north-
westward a considerable distance, the previously reported limit being the Uelle
River, Zaire. Like the Congo specimens of ignea, these birds have the rec-
trices brownish, only washed with rufous.

Family Paridae

Parus leucomelas guineensis Shelley. — One male, testes not enlarged, was
taken at the edge of the forest 8 June; weight 20 g; also seen 12 and 13 June.

Family Nectariniidae

Anthreptes longuemarei haussarum Neumann. — One male, 1 female,
gonads not enlarged, were taken at the edge of the forest 7 and 13 June; weight
male 14 g, female not recorded.

* Anthreptes rectirostris tephrolaema (Jardine and Fraser). — Five males,
gonads varying from small to large, were taken in trees at the forest edge, 12 to
16 June; weights 10 g in 4 cases, 11 g in the other.

Anthreptes collaris somereni Chapin. — One male, testes enlarged, 3
females, ovaries small, were collected in fruiting trees at the forest edge 5 to 17
June; weight of male 10 g, females 8 g each.

^Nectarinia seimundi traylori Wolters. — One specimen, sex undeter-
mined, was taken in a fruiting tree at the forest edge 15 June.

Nectarinia olivacea cephaelis (Bates). — This Sunbird must be common
in the Ouossi River area; 8 specimens, 3 males with large testes, 5 females with
small to slightly enlarged ovaries, were taken in the riverine forest 31 May to 8
June; weights males 11, 11.5 and 12 g, females 9 g in 1, 10 g in the other 4
individuals.

* Nectarinia cyanolaema octaviae Amadon. — Two males with large
testes were shot in fruiting trees at the forest edge 18 and 20 June; weights 15,

16 g.

Nectarinia superba superba (Shaw). — One male, testes large, was col-
lected in tree tops at the forest edge 17 June; weight 17.5 g.

Family Emberizidae

Emberiza cabanisi cabanisi (Reichenow). — One female, ovary not en-
larged, was taken at the forest edge 31 May; weight 22 g.

Family Estrildidae

Parmoptila woodhousei woodhousei Cassin. — One male, testes slightly
enlarged, was netted in swampy undergrowth of the forest 10 June; weight 9.5
g, and extends the known range of this subspecies eastward about 500 miles
along the northern limits of its range. In the Central African Republic it was

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No. 287

known earlier only from the extreme western area, bordering on Cameroon.

*Nigrita bicolor brunnescens Reichenow. — Two females, 1 with small
and 1 with enlarged ovary, were taken near the forest edge 10 and 12 June;
weights 13 and 15 g.

*Nigrita canicapilla canicapilla (Strickland). — Two females, 1 with en-
larged, 1 with small ovary, were collected 15 and 19 June at the forest edge;
weight 20 g each. The present birds are very slightly paler gray above than a
series of schistacea from western Uganda, but the two races must meet not far
to the east of the Ouossi River area. The birds had been eating small fruits,
probably of the trees from which they were collected.

Euschistospiza dybowskii (Oustalet). — One female with small ovary, and 1
bird of undetermined sex (male by plumage), were collected in dense thicket at
the forest edge 8 and 20 June; weight of male (?) 13 g, female 12 g; stomach
contents grass seeds and insects. These specimens fill a considerable blank in
the records shown by Hall and Moreau (1970: map 385).

*Mandingoa nitidula schlegeli (Sharpe). — One male, testes not enlarged,
was netted in forest undergrowth 12 June; weight 11 g.

Pyrenestes ostrinus ostrinus (Vieillot). — The Black-billed Seed-cracker
must be common in the Ouossi River area as 6 examples, all with small gonads,
were taken in mist nets in the swampy forest undergrowth 2 to 19 June; weights
males 21, 21.5 22 g, females 21, 23, 23 g. Mackworth-Praed and Grant
(1973:695-696) treat ostrinus and rothschildi as two sympatric species on the
assumption that “it seems very doubtful that large-billed and small-billed birds
would interbreed in any one locality ...” On the other side of the argument,
Chapin (1954:493) writes “that birds of differing sizes may mate with each
other cannot be doubted” and gives specific instances. It may be that before
man began to clear and otherwise intrude into previously uniformly forested
areas there may have been separation of the two “species” but the fact that
they do cross and that variants of all degrees of intermediate size occur to-
gether in many areas indicates that the birds do not “recognize” specific dif-
ferences. And they are, biologically, the reliable judges. The present 6 birds
show both ostrinus and rothschildi characters in bill size. A similar situation
prevails in a long series of specimens from western Uganda.

*Spermophaga ruficapilla ruficapilla (Shelley). — The Red-headed Blue-
bill must be very abundant in the Ouossi River area as 14 specimens, 8 males,
all with small or only slightly enlarged testes, and 6 females, all with small
ovaries, were taken, largely in mist nets, in the dense forest undergrowth 2 to
12 June; weights 22 to 27 g (average 24.3) in males, 24 to 28 g (average 24.8) in
females. The present records extend the known range of the species over a
hundred miles to the northwest.

Family Ploceidae

Ploceus nigricollis nigricollis (Vieillot). — One female, ovary slightly en-
larged, was collected in a fruiting tree at the forest edge 18 June; weight 31 g.

1978

The Lathrop Ornithological
Central African Expedition

21

Ploceus cucullatus bohndorffi Reichenow. — Two males, testes enlarged,
were taken at the forest edge 8 and 12 June; weights 41, 47 g.

*Malimbus nitens microrhynchus Reichenow. — Two females, ovaries
not or only slightly enlarged, were netted in riverine forest undergrowth 4 and
15 June; weights 30, 34 g. These birds are small, wings 80 and 82.5 mm, and
agree with a series from western Uganda. They extend the known range of
microrhynchus about 300 miles to the northwest.

Malimbus malimbicus crassirostris Hartert. — Three specimens, all in
non-breeding state, were taken 8 to 19 June; weight male 36 g, females 31 and
33 g. These birds are close to crassirostris from western Uganda and fill a
sizeable gap in the data recorded in Hall and Moreau (1970: map 348).

Family Dicruridae

Dicrurus ludwigii sharpei Oustalet. — The Square-tailed Drongo must be
fairly common in the Ouossi River forests, as 5 specimens were taken, 3 males
with enlarged or slightly enlarged testes, 2 females with small ovaries, 3 to 11
June; weights males 31, 31, 32 g, females 26, 30 g.

*Dicrurus atripennis Swainson. — One female, ovary slightly enlarged,
was netted in open forest undergrowth 14 June; weight 42 g; stomach contents
mainly beetles.

ACKNOWLEDGMENTS

The Lathrop Central African Republic Expedition of 1976 was made pos-
sible by the generosity and interest of Mr. and Mrs. Charles F. Lathrop of
Ellensburg, Washington. Not only are our thanks due to the Lathrops for their
generous sponsorship, but also to Andrew Williams and his African assistants
for their excellent work in obtaining the present collection and the new knowl-
edge its study has revealed, and to Mrs. Reese Hale Taylor who kindly typed the
manuscript.

LITERATURE CITED

Anonymous (probably J. SABATER PI). 1963. El misteriosa “pajaro indicador de cola
de lira.” Zoo: Revista de Parque Zoologico de Barcelona, no. 2:43.

Archer, A.L. and R.M. Glen. 1969. Observations on the behavior of two species of
honey-guides, Indicator variegatus (Lesson) and Indicator exilis (Cassin). Los
Angeles County Museum Contr. Sci. no. 160:1-6.

Beatty, H.A. 1963. The lyre-tailed honey-guide in the Ivory Coast. Bull. British Or-
nithologists’ Club, 83:100-101.

Chapin, J.P. 1932. The birds of the Belgian Congo. Pt. I. American Museum Nat. Hist.
Bull. 65:1-756.

1939. The birds of the Belgian Congo. Pt. II. American Museum Nat. Hist.

Bull. 75:1-632.

22

Contributions in Science

No. 287

. 1953. The birds of the Belgian Congo. Pt. III. American Museum Nat. Hist.

Bull. 75A:1-821.

. 1954. The birds of the Belgian Congo. Pt. IV. American Museum Nat. Hist.

Bull. 75B: 1-846.

. 1962. Sibling species of small African honey-guides. Ibis, 104:40-44.

Clancey, P.A. 1970. Miscellaneous taxonomic notes on African birds. XXIX. Durban
Museum Novitates 8, pt. 20:375-390.

. 1972. A catalogue of birds of the South African Sub-Region. Supplement no.

2. Durban Museum Novitates 9, pt. 12:163-200.

Friedmann, H. 1955. The Honey-guides. United States National Museum Bull. 208:1-
292.

. 1968. Zenker’s Honey-guide. Journal fur Ornithologie, 109:276-283.

. 1969. The status and distribution in Uganda of the white-bellied kingfisher,

Alcedo leucogaster leopoldi. Los Angeles County Mus. Contr. Sci. no. 158:1-6.
. 1970. Phenotypic potential and speciation in Indicator and Prodotiscus. Os-
trich, Supplement, 8:21-26.

Friedmann, H. and J.G. Williams. 1968. Notable records of rare or little-known birds
from western Uganda. Revue Zoologique et Botanique Africain, 77, nos. 1-2:1 1-36.

. 1971. The birds of the lowlands of Bwamba, Toro Province, Uganda. Los

Angeles County Mus. Contr. Sci. no. 211:1-70.

Hall, B.P. and R.E. Moreau. 1970. An atlas of speciation in African passerine birds.
British Museum (Nat. Hist.): xv+ 1-423.

Irwin, M.P.S. and C.W. Benson. 1966. Notes on the birds of Zambia. Pt. II. Arnoldia,

2, no. 37:1-21.

Keith, S., A. Twomey, H. Friedmann, and J.G. Williams. 1969. The avifauna of the
Impenetrable Forest, Uganda. American Museum Novitates no. 2389:1-41.
Mackworth-Praed, C.W. and C.H.B. Grant. 1970-1973. Birds of west-central and
western Africa. 2 vols. 1-670; 1-818.

Malbrandt, R. 1952. Faune du Centre Africain Francais (mammiferes et oiseaux). ed.

2. in Encyclopedic biologique. Paris, 15:1-616.

Malbrandt, R. and A.R. Maclatchy. 1949. Faune L’Equateur Africain Francais, 1.

Oiseaux. in Encyclopedic biologique, Paris 35:1-460.

Meyer de Schauensee, R. 1949. Results of the Carpenter African Expedition 1947-1948.

Notulae Naturae, Academy Natural Sci. Philadelphia, no. 219:1-16.

Morony, John J. Jr., W.J. Bock, and J. Farrand, Jr. 1975. Reference List of the Birds of
the World. American Museum Nat. Hist. 1-207.

Prigogine, A. 1971. Les oiseaux de L’ltombwe et son hinterland. Musee Royal de
l’Afrique Centrale,-Tervuren, Annals, ser. in 8° Ser. Zool., no. 185:1-298.
Schouteden, H. 1962. La faune ornithologique des districts de la Mongala et de
l’Ubangi. Musee Royal de l’Afrique Centrale,-Tervuren. Documentation
Zoologique no. 3:1-144.

. 1963. La faune ornithologique des district de 1’Ituri. Musee Royal de V Afrique

Centrale,-Tervuren. Documentation Zoologique no. 5:1-144.

White, C.M.N. 1965. A revised check-list of African non-passerine birds. Lusaka, Govt.
Printer: 1-299.

Williams, J.G. and H. Friedmann. 1965. The pygmy honey-guide. Indicator pumilio
Chapin, in East Africa. Bull. British Ornithologists’ Club, 85:21-22.

Zimmerman, D. A. 1972. The avifauna of the Kakamega Forest, western Kenya, includ-
ing a bird population study. American Museum Nat. Hist. Bull. 149, art. 3:259-339,.

Accepted for publication January 19, 1977.

501, 73

NUMBER 288
MARCH 17, 1978

FOLIICOLOUS ASCOMYCETES 1:
THE CAPNODIACEOUS GENUS SCORIAS,

REPRODUCTION

By Don R. Reynolds

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCENCC

Published by the NATURAL HISTORY MUSEUM
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CONTRIBUTIONS IN SCIENCE contain articles in the earth and life sciences, presenting
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Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

FOLIICOLOUS ASCOMYCETES 1:

THE CAPNODIACEOUS GENUS SCORIAS
REPRODUCTION1

By Don R. Reynolds2

Abstract: Scorias spongiosa (Schw.) Fries development is analyzed by
use of sequential collections from Georgia. The soma develops during the fall to
winter season. The morphology of the anamorphic centrum remains unchanged
as the soma matures. The teliomorphic centrum is dothideaceous; a sterile
element in this centrum is interpreted as laterally positioned periphyses; the
ascus is functionally bitunicate. A protocol to substantiate definite proof of
reproductive state pleomorphism is discussed.

INTRODUCTION

Scorias Fries, a classic North American representative of sooty mold As-
comycete fungi, is known in the earliest American mycological literature. This
capnodiaceous genus is based on an early known North American species, S.
spongiosa (Schw.) Fries. This fungus occurs in a typical sooty mold plant-
surface habitat in saprobic association with insect exudate. Although the
distribution of the fungus is known to be along the U.S. east coast from northern
Florida to Maine, S. spongiosa has received little attention since its initial
impact on mycological literature of the 19th century.

The labyrinthic stroma formed by the dark mycelial growth, preceding
pseudothecium formation, was first termed “sponge.” Initially Schweinitz
( 1 822) and later Fries ( 1 829) in the type description of the genus, commented on
the waxlike appearance of the mycelium and the change in the “sponge”
matrix from brittle to soft upon absorption of moisture. The nam e Scorias was
utilized by Fries to denote the stromatal resemblance to slag as the structure
developed on leaves and branches of the American beech, Fagus grandifolia
L. S. spongiosa was illustrated by Ellis and Everhart (1892) with a drawing
showing ascocarps and pycnidia (as “spermogonia”) originating from the same

Review Committee for this Contribution
Christopher Davidson
M.L. Farr
James Kimbrough

2Senior Curator of Botany, Natural History Museum of Los Angeles County, 900
Exposition Boulevard, Los Angeles, California, 90007. USA

2

Contributions in Science

No. 288

“fertile branch.” They placed the monotypic genus in a taxon, the Peris-
poriaceae, defined with “ostiolum obscure or wanting . . . opening irregu-
larly.” Von Hoehnel (1910) described the hyphae as copious, partially slimy
and fused together bundlelike. He characterized (1909) the pseudothecium
centrum as cartilaginous-gelatinous due to the nature of the component sterile
elements. Ascus dispersal was attributed to an apical swelling of the centrum.
“Ein ausgesprochenes Ostiolum fehlt stets.” Few other morphological insights
can be gleamed from subsequent literature (i.e., Lloyd 1921; Batista & Ciferri
1963b).

This paper concerns the developmental morphology of Scorias spongiosa.
A taxonomic monograph of the genus Scorias Fries is to appear later as an
additional contribution from ongoing studies focusing on foliicolous fungi.

MATERIALS AND METHODS:

The sequential collections selected for detailed analysis represent periodic
sampling from Athens, Georgia — i.e. LAM 200001 (25 X 1971), LAM 200004
(12 XI 1971), LAM 200006 (02 XII 1971), LAM 200007 (20 XII 1971). Collec-
tion LAM 200008 (21 I 1972) was utilized for illustration of mature material;
information from collection LAM 200000 (18 VIII 1973) was incorporated for
data on early development. Additional specimens examined from Georgia in-
cluded LAM 200002 (02 II 1972), LAM 200004 (09 II 1972), LAM 200005 (20 II
1972), LAM 200009 (05 IV 1972), LAM 200010 (24 IV 1970), LAM 200011 (11
V 1972), LAM 200012 (11 V 1970) and LAM 200013 (10 V 1973). Other her-
barium specimens examined included those from BPI (Brown VII 1932, Shear
4 X 1935, Shear 1 II 1903); CUP (Howard Cayuga Flora 48), FLAS (1862,
1863, 1864, 21 183, 46426); ILL (Rogers 14-15 III 1958), Ellis, North American
Flora (1363 a & b), Ravanel Fungi Americana (334 and 1877). Schweinitz
collections from herbaria E, FH, K, PH and UPS were additional non-
Georgian specimens examined.

Fresh material and 2% KOH revived dry material were cut on a Bailey
freezing microtome and mounted in lactophenol or lactophenol-cotton blue;
additional material was killed and fixed in FAA, paraffin embedded and the
resulting sections stained with hematoxylin and methylene blue.

Camera lucida drawings were made by DRR; composite drawings from
direct observation of sectioned material were done by F.E. Runyan.

RESULTS

Thallus Development

The general appearance of early thallus formation was as cream to buff
colored tufts of mycelium on twigs and on some leaves of the American beech
(LAM 200000). The fungus soma developed in accumulations of the exudate of
the wooly aphid insect, which utilizes F. grandifolia as a host plant. Limited
mycelial growth was seen on the ground at time of collection. The nonpig-

1978

Foliicolous Genus Scorias

3

mented hyphae exhibited two distinct zones in the wall. In cross section of
older hyphae (Fig. 1 A), the outermost zone was widest, measuring an average
15 fjL. The outer boundary of the inner- wall zone was sharply delineated and
measured approximately \/x in diameter. The cell lumen containing the proto-
plast averaged 6/jl in diameter. Cells were generally rectangular; size was
variable (Fig. IB). The outer-wall zone was present in the apical cell of
presumed actively growing hyphae (Fig. 1C) and became wider in diameter than
the inner-wall zone in cells progressively subtending the apical cell. The wall
formed a constriction at sites corresponding to the location of septa traversing
the cell lumen. In median cell focus, a centrally located gap could be detected in
the isodiametric septa.

Figure 1. Hyphae. A. Cross section of hyphae, approx. 1300X; B. hyphae illustrating
two wall zones, approx. 1000X; C. tip of young hypha, approx. 1000X.

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Contributions in Science

No. 288

Hyphal organization was effected largely by adherence of outer wall sur-
faces; reinforcement by cell fusions resulted in cytoplasmic continuity. Hyphae
adhering together in small numbers for short distances, diverging individually
or in small numbers and readhering with single or grouped hyphae, maintained
a loose mycelial subiculum in localized areas on the supporting plant surface.
The basic hyphal organization appeared as strands. Strand construction was
similar throughout thallus development. The strand was usually rounded in a
cross-section outline; the component hyphae were more or less parallel.

The somatic mycelium was well developed at a point in time represented
by specimen LAM 200001. The hyphal wall construction was similar to that
described in LAM 200000. The outer-wall zone measured 5-15 /x in diameter. An
irregular verrucose deposition could be detected on hyphal surfaces in direct
contact with other hyphal strands which were exposed to the atmosphere. The
inner-wall zone measured approximately 1 /jl in diameter. An interzonal area
was identifiable which varied in width from almost nil to \/jl: it could be
recognized by dark lines of varying length, oriented generally perpendicular to
the axis of the hyphae, radiating from the outer surface of the inner-wall zone
into the outer-wall zone. Pigmentation occurred first on exposed strand sur-
faces. The inner-wall zone of mature hyphal strands was initially achromatic and
became darkened with a deposition of a pigment assumed to be melanin. The
outer-wall zone was also initially achromatic and became darkened with a
pigment similar to that of the inner zone, but of less intensity.

Mature stromata produced by the somatic mycelium of S. spongiosa had a
definite internal structure (Figs. 2 & 3). Collection LAM 200013 is representa-
tive of mature stroma structure. It measured 1 1 .6 cm in length and 5.5 cm at its
highest point. Fig 2 is a diagrammatic representation of the stroma viewed
lengthwise; Fig 3 represents a cross section at a level indicated by the arrow in
Fig 2. The mycelial development radiated upward and outward from the sup-
porting twig. The hyphae coalesced to form mycelial strands, which were

1

Figure 2. Diagrammatic representation of stroma lengthwise profile. Note enlargement
of stroma in area represented by arrow in Figure 3.

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Foliicolous Genus Scorias

5

Figure 3. Cross-section view of mature stroma, approx. 3X. Supporting twig would be
located at lower center portion.

largest in the lower center of the stroma. These central strands were formed in
a lacunate region within the stroma which was also the highest portion (Arrow,
Fig. 2; Fig. 3). Above the central labyrinth, the mycelial strands branched
more frequently resulting in a compacted layer 1-2 cm deep, which formed the
outermost region of the stroma. Reproductive structures ultimately developed
on the exposed surface of the stroma. In the outermost compacted region of
developing stroma, the hyphae comprising the outer layers of the component
mycelial strands were fully pigmented; hyphal walls in the inner mycelial
strand core were hyaline to yellowish. In mature stroma (Fig. 3) collected in
late spring and bearing pseudothecia (LAM 200013), the strands in the outer-
most compacted portion were fully pigmented and most were surrounded by a
nonpigmented layer representing the individual hyphae of new growth on the
strand surface.

The dry stroma enlarged on contact with moisture. A rough estimation
was made of structure modification upon wetting by calculating weight in-
crease. Sections cut of collection LAM 200013 were weighed before and after
wetting in a 10% formalin solution. An average 1.4% increase in weight was
noted. Lack of appropriate material precluded confirmation of these data by
use of additional stroma.

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Contributions in Science

No. 288

CONIDIOGENY

A phialidic anamorphic centrum was present at the tips of the mycelial
strands in collections LAM 200000 and LAM 200014 on the cream to buff-
colored tufts of mycelium (Fig. 4). Within the hyaline strand apex, the con-
idiogenous system was initiated by profuse transverse cell division in the
centrally positioned parallel hyphae.

The short cells of the centrally located dividing hyphal strands were dis-
tinct from larger elongate cells of the hyphae in the outermost strand layers.
Additional hyaline cells were produced from these short cells, which in turn
branched toward the strand center. Sympodial branching was initiated from a
site immediately below the apical septum and generally opposite a conidioge-
nous site (Fig. 8B). Limited apical expansion resulted in a branched system of
phialogenic hyphae of one to several cells in length. Phialospores were pro-
duced from usually only one place on the conidiogenous cell near the upper-
most septum, less frequently in the middle of the cell, or at the hyphal tip ( Fig.
8A-E). No observations were made on initial spore development from living
material. The first conidium was apparently formed as a small tubular protru-
sion which was blown out from the conidiogenous cell (Fig. 8 A). Successive
spores were produced through a collarette (Fig. 8D, E).

A dark brown pigment appeared in the inner-wall zone of the short-celled
hyphae from which the conidiogenous centers were initiated, as well as in cells
of adjacent strand hyphae with longer cells. The outermost nonpigmented
hyphae of the strand apex was separated from the enlarged conidiogenous
centers from this time on. Lateral expansion of the anamorphic centrum
coincided with proliferation of conidiogenous hyphae and production of conidia;
the conidia became compacted in the middle of the conidiogenous center,
creating a locule. A necklike extension was developed beyond the sporogenous
area by a convergence and elongation of a layer of dark strand hyphae surround-
ing a central cylinder of nonbranching sterile hyaline phialogenic hyphae. The
outer darkly pigmented hyphal layer eventually ceased to grow; the hyaline
inner layer of hyphae extended beyond the outer layer, became reflexed,
resulting in what Batista and Ciferri (1963a) referred to as a fimbriated ostiole.
Conidia appeared to be initially forced through a space between the hyphae of
the hyaline core, thus creating a channel to the end of the neckline extension.
These and subsequent conidia were exuded in a slimelike matrix which collected
as a droplet at the apex of the neck.

The phialidic anamorphic centrum was present on the surface of the
well-developed stroma; it was well represented in October collections as well as
in collections from later in the year. Initiation of these conidium forming areas is
identical to that described from collections LAM 200000 (Fig. 5); the resulting
conidia were identical. Flowever, the phialidic anamorphic centrum was formed
in short mycelial strands of darkly pigmented hyphae which radiated from the
stroma surface. The length of the subtending strand varied. None of these
pycnidium-like structures was seen in this later stage to have an apical continua-
tion of the outer mycelial strand hyphae into a necklike extension.

1978

Foliicolous Genus Scorias

7

Figure 4. Asexual fruit body with phialidic anamorphic centrum, approx. 300X.

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Contributions in Science

No. 288

Figure 5. Asexual fruit body with phialidic anamorphic centrum, approx. 1000X.

Figure 6. Stroma surface detail illustrating ascocarps, approx. 30X.

Ascogeny

Ascocarp formation was initiated by internal proliferation within a myce-
lial strand found on the stromatal surface (Fig. 6) and culminated in a prehyme-
nial structure designated here the protopseudothecium. Eight or more indi-
vidual hyphal strands from the surface network recoalesced in strand forma-

1978

Foliicolous Genus Scorias

9

tion. Elongation of the strand appeared to be from the apical ends of the
component hyphae. The mycelial strand organization was initially similar to
that found in the central and compacted areas of the stroma, whereby parallel
anastomosing hyphae formed into a rounded branch; the relative position of
hyphae in the outer to central areas remained constant. All cells were found to
be melanin-like pigmented. The onset of protopseudothecial differentiation
was signaled by three areas of hyphal proliferation. A mycelial strand where a
protopseudothecium was to be initiated was comprised of several hyphal
zones; configuration patterns could be discerned in the outermost hyphal layers,
in the middle hyphal layers and in the remaining innermost hyphae. The
outermost layers did not depart from the basic parallel strand pattern (Fig. 7B).
The hyphae in the middle layers began three-dimensional dichotomous and
trichotomous branching. The cells were shorter but similar in cell shapes as
compared to the outer most parallel hyphae (Fig. 7D). The hyphae in the
innermost layers proliferated by formation of small isodiametric hyphae. These
frequently branched, intermingled hyphae were generally oriented inwardly,
perpendicular to the strand axis (Fig. 7E). All hyphae in the proto-
pseudothecium had the basic 2-zone wall construction described earlier. Pig-
mentation developed in the outer layers containing the parallel, and the di- and
trichotomously branched hyphae (Fig. 7B, C), but not in the isodiametric
hyphae in the center of the protopseudothecial strand (Fig. 7D, E). Develop-
ment of the outer layers of protopseudothecial strand kept pace with innermost
core layer so that the pigment containing layers surrounded the inner core of
hyaline isodiametric hyphae at all times. Proliferation of the latter and sub-
sequent increase in width began in the lower portion of the proto-
pseudothecium and became widest in the center of the ascocarp and terminated
in an ostiole at the uppermost part.

Thus, the basic ascocarp subtended by a stalk (Fig. 7 A) was formed before
appearance of the ascogenous system. No further strand development ap-
peared to occur after the inner core of hyaline hyphae was produced in the
protopseudothecium. The portion of the protopseudothecial strand forming the
ascocarp wall appeared plectenchyma-like in that the intertwined cells became
affixed together, presumably by wall fusion. However, the centrally located
nonpigmented hyphae retained their individual identity (Fig. 7E). In a longitu-
dinal section of a protopseudothecium, the tissue might appear as
pseudoparenchyma in both the wall and the central mycelial mass. This effect,
as represented in the drawings such as those by Batista and Ciferri (1963b) for
S. spongiosa, is due to the intermingled, highly-branched hyphae in the pig-
mented and nonpigmented layer in section showing the lumen of component
cells from various angles separated by the wide walls.

The ascogenous system (Fig. 7F) appeared at the lower end of the
pseudothecium. No ascogonium was detected; asci appeared to originate as
croziers on proliferating ascogenous hyphae. Asci matured first in the center of
the developing hymenium. The ascus wall was isodiametrically thickened only
until the ascus mother cell elongated to full size. An ascus devoid of spores was
cylindrical (Fig. 8F). The elongating ascus mother cells appeared to push up-

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No. 288

Figure 7. Ascocarp, approx. 1000X. A. mycelial strand subtending ascogenous center as
stalk; B. base of asocarp; C. dark tissue layer delimiting ascocarp; D. inner zone of
hyaline hyphae; E. lateral periphyses lining locule; F. ascogenous hyphae bearing asci;
G. ostiole.

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Foliicolous Genus Scorias

11

ward among the nonpigmented hyphae occupying the center of the
pseudothecium (Fig. 7). As the hymenium developed, the nonpigmented
hyphae were displaced from the lower 75 percent of the original areas they
occupied. The interface at the place of origin of nonpigmented hyphae with
pigmented strand hyphae, effectively limited displacement. The resulting dis-
tortion created space within the pseudothecium (Fig. 7).

The shape of an ascus containing spores had been modified to obpyriform;
the wall of such an ascus was thickened at the ascus apex with an evident nasse
apicale (Reynolds 1971) and tapered in width toward the ascus base (Fig. 8G).
Mature ascospores were hyaline and traversed by three septa. Discharged asci
were found to have a discernible outer tunica surrounding a tubular inner
tunica. Ascospores were frequently seen attached to the outer surfaces of
ascocarps, some having germinated. No ascus was found to be discharged
intact from the ascocarp as suggested by Ellis and Everhart (1892), although in
some preparations, pressure applied to a coverslip would result in dislocation
of individual asci and whole hymenia through the ostiole.

Figure 8. Phialides and asci, approx. 2000X. A, B. enterblastic monophialidic con-
idiophores bearing young conidia; D, E. phialides with collarettes; C. conidia; F. young
ascus before ascosporogenesis; G. mature spore-bearing ascus.

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No. 288

DISCUSSION

The terms teliomorphosis and anamorphosis have been introduced (Hen-
nebert and Weresub 1977) in relation to a restatement of the International Code
of Botanical Nomenclature, Article 59, “Names of fungi with a pleomorphic
life cycle and of fossils assigned to form genera.” These terms refer to basic
reproductive modes of the holomorphic fungus and were defined as follows:
teliomorphosis — “characterizing form that is involved in producing meiotic
diaspores — in Ascomycetes, an ascocarp or its equivalent producing asci and
ascospores. . . anamorphosis — “The imperfect state . . . the asexual mi-
totic diaspore expression of a fungus ...”

The term anamorphic centrum is utilized here as a Deuteromycete applica-
tion of the Luttrellian Ascomycete centrum concept This and other related terms
are defined as follows: teliomorphic centrum = centrum sensu Luttrell (1951);
anamorphic centrum = the reproductive or conidiogenous system component of
an asexual fruit body; phialidic anamorphic centrum = an anamorphic centrum
where the conidium is enteroblastically produced via a phialide.

The conidiogenous system described here consists of enteroblastic, ba-
sically monophialidic, conidiophores resembling that of two epimycotic
Coelomycete genera. The illustrations by Seeler (1943) of Eleuthoromycella
mycophyla von Hoehnel and especially Eleutheromyces sublatus (Fr.)Fuckel
indicate strikingly similar acropleurogenous branched conidiophores. Both
species are described as having wall tissue which is pseudoparenchymatous
(textura angularia) rather than of parallel, somewhat gelatinized hyphae (tex-
tura oblita), as found in S. spongiosa. The pigmentation of E. subulatus is a
nectriaceous burnt sienna, and that of E. mycophyla is carbonaceous. The walls
of S. spongiosa pycnidia range from hyaline to melinoid.

Only one anamorphosis is associated with the life history of S. spongiosa.
The term anamorphic centrum has been used to emphasize a continuity in
terms of the conidiogenous system and the conidia produced in association with
the stroma in all stages of development. Morphological forms of the asexual
fruit body found here can be assigned to various pycnidial genera. Following
Batista and Ciferri (1963a), these would be Leptoxyphium (globose, sessile, with
no neck), Podoxyphium (globose stalked, with no neck) and Microxyphium
(cylindrical, with a neck). Hughes (1976) speculatively refers to similar mor-
phological forms as Polychaeton and Conidiocarpus (and perhaps
Scolecoxyphium). The position of the anamorphic centrum in the rapidly
expanding and rounding mycelial strands (i.e., LAM 200000) would seemingly
cause a cylindrical pycnidium-like structure. Continued growth of the sur-
rounding hyphae would produce a “neck.” This observation is made especially
clear when conidiogenous systems are to be found centrally located in mycelial
strands with the ostiole eliminated by the continued growth of the strand tip.
The conidiogenous systems found in the more mature stroma develop at the
ends of narrow hyphal strands. Unlike the pycnidia with a “neck,” these
strands extend from the surface of a stroma and terminate with the formation
of the conidiogenous center.

1978

Foliicolous Genus Scorias

13

Therefore, the generic concepts of these pycnidiaceous fungi as set forth
in literature such as Batista and Ciferri (1963a) and Hughes (1976) are ques-
tionable; many of these appear to be based on growth cycle artifacts given
undue taxonomic significance. Experimental data derived from pure culture
work would go far in resolving the problem. Detailed observations on a
chronologically obtained series of collections from a single natural locality,
such as utilized in this study, may prove a valid alternative.

The teliomorphic type is modified dothideaceous. The ascus produced by
S. spongiosa is functionally bitunicate; microscopic mounts reveal the nasse
apicale apparatus proposed by Chadefaud and explained by Reynolds (1971).
The sterile elements peripherally produced from the wall, which hang into the
locule, are not pseudoparaphyses sensu Wehmeyer. Therefore, in agreement
with von Arx and Muller (1975), the concept of these structures as pleosporace-
ous(Corlett 1973), is rejected. These structures originate as outgrowths from the
pseudoparenchymatous peripheral layer of the ascocarp locule and the ostiole.
The term for these structures in an ostiole is periphyses. Because they are
produced to a lower level in the ascocarp but are not attached in the hymenium
as are paraphyses and additionally are not involved in centrum formation as are
pseudoparaphyses, they are regarded as laterally positioned periphyses or
periphysoids (Barr 1976). Samuels (1973) discussed centripetal paraphyses and
apical paraphyses as found in unitunicate species.

In Scorias, as in other Ascomycetes, taxon definition is typed with the
teliomorphosis. No Ascomycete species is known to produce more than one
teliomorphosis sexual state, but many are known which produce one or more
anamorphoses from the same mycelium or thallus. My attempts to work with
Scorias spongiosa in artificial culture failed to yield the desired reproductive
states. Instead, a hopefully acceptable compromise has been utilized involving
sequentially collected herbarium specimens, which is similar to that utilized by
Luttrell and Muthappa (1974). Holomorphic data was derived from collections
taken periodically from a particular locality over a time span during which the
fungus developed in nature. Additional and supportive information was derived
from usual herbarium specimens available on loan from the institutions previ-
ously mentioned. Consequently, the author is reasonably convinced, in absence
of definite proof sustained by data from artificial culture methods, that the
stroma development is accurately presented for a localized strain of S. spon-
giosa, and that the reproductive states are likely to be those actually produced
by the species, rather than being associated with the stroma of S. spongiosa by
virtue of habitat preference of sooty mold fungi with similar morphological
features.

Imperative in profiling the capnodiaceous whole-organism is an absolute
certainty that the alternate reproductive states are biologically associated. M til-
ler (1971), in a review of perfect-imperfect connections in Ascomycetes,
pointed out that use of pure culture techniques has allowed definite proof of
connections between the teliomorphosis and any anamorphoses produced by
one organism. Biological connection of alternate states are definitely proved in

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No. 288

artificial cultivation, where this method is possible, by use of spore-to-spore
cultivation, or by germination of one spore form to give rise to a mycelium in
culture which produces an alternate state. Definite proof of biological connec-
tion is lacking, and only suspected, where demonstration of a single hyphal
strand organically connecting several alternate states is found in randomly
obtained herbarium specimens.

My concept of sooty mold fungi is derived from several thousand collec-
tions I have made in all of the neotropical countries and in some areas of the
paleotropics, on the examination of all herbarium specimens I can locate which
have been cited in the literature, and from attempts at experimental culture
work with field-derived isolates. Consequently, I strongly advocate cogni-
zance of the specialized microenvironment into which sooty mold fungi have
adapted when systematic judgments are formulated as well as in the perception
of pleomorphic or alternate reproductive state relationships. Therefore, espe-
cially in capnodiaceous fungi, systematic relationships should be determined
from data derived from artificial culture of the species as well as from speci-
mens collected in nature. Analytical analysis of the holomorphic sooty mold
species demands a nonintuitive laboratory protocol in order that realistic
taxonomic innovation can be made.

The intuitive “analytical analysis” of the interpretation of the sooty mold
fungi was summarized by Hughes (1976). This viewpoint is in direct contrast to
the rendering of systematic judgment which incorporates experimental data
derived from a protocol utilizing pure culture techniques such as that demon-
strated by Simmons (1969), or a chronologically obtained series of specimens
from a single natural collection locality such as that utilized by Luttrell and
Muthappa (1974). Hughes advocated prima facie establishment of alternate
reproductive state association in the sooty mold colony as may be found in
randomly collected herbarium collections. The a priori taxonomy proposed by
Hughes (1976) contains the highly interesting, but biased, insight of a Fungi
Imperfecti specialist. However, his philosophical commentary was unfortu-
nately expressed in nomenclatoral jargon and, lacking experimental data, falls
short of proper systematic documentation.

The annual growth and reproduction cycle detected in the sooty mold
fungus S. spongiosa is regarded as indicative of efficient utilization of available
microenvironmental resources under the influence of overall seasonal changes.
This species and others in the family Capnodiaceae von Hoehnel exhibit mor-
phological and life-history features, which would give advantages in coping
with the stressful plant surface-atmosphere environment in which they exist.
Thus, concepts of thallus and fruit body development should be justifiably
influenced by habitat-related data. The sooty mold colony produced in nature
by the capnodiaceous fungi should basically be regarded as an environmentally
influenced association of species (Reynolds 1975; Reynolds and Pohlad 1974).
Any attempt at analytical analysis of the fungi in the plant surface microenvi-
ronment will have to be based on a convincing, nonintuitive demonstration of

1978

Foliicolous Genus Scorias

15

biological relationships between teliomorpheses and anamorpheses of fungi
which thrive in this habitat. Definite proof of these relationships is necessary
because ultimately any patterns of reproductive state associations should be
reflected in the taxonomy of these highly evolved Ascomycetes.

ACKNOWLEDGMENTS

Acknowledgment is made to Dr. E.S. Luttrell for assistance with the
Georgia collections, and to the National Science Foundation, The American
Philosophical Society, the Los Angeles County Natural History Museum
Foundation for financial support.

LITERATURE CITED

Arx, J. A. von, andE. Muller. A reevaluation ofthe bitunicate Ascomycetes with keys
to families and genera. Studies in Mycology No. 9. Centraalbureau voor Schimmel-
cultures Baam, The Netherlands. 159 p.

Barr, M.E. 1976. Perspectives in the Ascomycotina. Mem. New York Bot. Garden.
28:1-8.

Batista, C., and R. Ciferri. 1963a. The sooty-molds of the family Asbolisiaceae.
Quademo 31:1-229.

and . 1963b. Capnodiales. Saccardoa 2:1-296.

Corlett, M. 1973. Observations and comments on the Pleospora centrum type. Nova
Hedwigia 24:347-360.

Ellis, J.B., and B.M. Everhart. 1892. The North American Pyrenomycetes. Publ. by
the authors. Newfield, New Jersey. 793 p.

Hennebert, G. L. and L. K. Weresub. 1977. Terms for states and forms of fungi, their
names and types. Mycotoxon 6:207-21 1.

Fries, E. 1829. Systema mycologicum 3:291.

Hoehnel, F von. 1909. Fragmente zur mykologie. VIII. Mitteilung, no. 354-406. Set-
zungsber. Kaiserl. Akad. Wiss., Wien, Math.-naturwiss. Kl., Abt., 1, 118:1157-
1246.

1910. Fragmente zur mykologie. XI. Mitteilung, no. 527-573. Setzungsber.

Kaiserl. Akad. Wiss., Wien, Math.-naturwiss. Kl., Abt. 1, 119:617-679.

Hughes, S.J. 1976. Sooty Moulds. Mycologia 68:693-820.

Lloyd, C.G. 1921. Mycological notes no. 65. C.G. Lloyd 6:1067.

Luttrell, E.S. 1951. Lniv. of Missouri Studies. No. 3, Taxonomy of the Pyrenomy-
cetes. 24:1-120.

and B.N. Muthappa. 1974. Morphology of a new species of Aulacostroma

(Hemisphaeriales). Mycologia 66:563-579.

Muller, E. 1971. Imperfect-perfect connections in Ascomycetes. In Taxonomy of
Fungi Imperfecti. B. Kendrick, ed. Univ. Toronto Press. 309 p.

Reynolds, D.R. 1971. Wall structure of a bitunicate ascus. Planta 98:244-257.

1975. Observations on growth forms of sooty mold fungi. Nova Hedwigia

26:179-193.

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No. 288

and B. R. Pohlad. 1974. Cyclic patterns of central Florida sooty molds. Proc.

Iowa Acad. Sci. 81:12-13.

Samuels, G.J. 1973. Perithecial development in Hypomyces aurantius. Amer. Jour. Bot.
60:268-276.

Schweinitz, L.D. 1822. Synopsis fungorum carolinae superioris. Schr. fung. Naturf.
Gesell. Lepizig 1:20-131.

Seeler, E.V. 1943. Several fungicolous fungi. Farlowia 1:119-133.

Simmons, E.G. 1969. Perfect states of Stemphylium. Mycologia 61:1-26.

Accepted for publication February 2, 1977.

50 7, 73

Cadges

NUMBER 289
MARCH 20, 1978

OTOLITHS AND OTHER FISH REMAINS
FROM THE CHUMASH MIDDEN AT RINCON POINT (SBa-1)
SANTA BARBARA-VENTURA COUNTIES, CALIFORNIA

By Richard W. Huddleston and Lloyd W. Barker

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCENCC

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OTOLITHS AND OTHER FISH REMAINS
FROM THE CHUMASH MIDDEN AT RINCON POINT (SBa-1),
SANTA BARBARA-VENTURA COUNTIES, CALIFORNIA1

By Richard W. Huddleston2 and Lloyd W. Barker3

Abstract: Material from a Chumash village site yielded 45 species of
fishes (29 kinds of bony fishes, and 16 kinds of sharks, skates and rays). The
fish remains, ranging in size from less than 1 to 75 mm, consisted of otoliths,
teeth, jaws (dentary, premaxillary and palatines), vertebrae, stings, dermal
denticles, dorsal spines and scales. As indicated by the species recovered, the
Chumash inhabitants fished from surface regions and surf zones to depths
exceeding 60 feet, utilizing hook and line, gill nets, traps, beach seines, har-
poons, spears and capture by hand. The most abundant identified remains
were otoliths of Genyonemus lineatus, scales of Sard inops caeruleus, and
dermal denticles of Squatina californica. None of the identified remains
showed signs of having been altered or modified for use in ornamentation.
Comparison of the fishing habits of the inhabitants of SBa-1 and Ven-3 in
Ventura indicated a similar level of advancement, which was superior and
more fishery-orientated than Ora- 190 in Orange County and SLO-2 in San
Luis Obispo County.

INTRODUCTION

Prior to the completion of this study Lloyd W. Barker lost his life acting as
an observer for the California Department of Fish and Game aboard a com-
mercial sealion capturing vessel, which capsized in the Santa Barbara channel.
Because of his untimely death, any errors or omissions are the responsibility of
RWH.

Fish and other faunal remains of midden sites usually have appeared in
archaeological literature as simple lists of identifications without any interpre-
tation or significance. In particular, remains of fishes have been neglected,
often being identified only as “fish” or “shark”. Otoliths (fish earstones) play
an important role in the identification of fishes recovered from middens. The
use of otoliths in archaeological work is not new, being employed first in the

1 Review Committee for this Contribution
John E. Fitch
W.I. Follett
Robert J. Lavenberg

2Section of Vertebrate Paleontology, Natural History Museum of Los Angeles County,
900 Exposition Boulevard, Los Angeles, California 90007.

3 Deceased.

2

No. 289

Contributions in Science

latter part of the nineteenth century (Ihering 1891). It was not until 1969,
however, that otoliths were used on a large scale (Fitch 1969a). Fitch noted
approximately 19 species of fishes represented by otoliths from a midden in
Ventura County.

All bony fishes (fishes other than sharks, skates, rays and chimaeroids)
possess some type of discrete aragonitic concretion in the semicircular canals
of the otic capsules, called otoliths. Although not well understood, the otolith
aids in the hearing and balance of the fish. Since almost all species of fishes
have distinctive otoliths, their specific identifications can be made with ade-
quate comparative material.

Bony fishes possess three pairs of otoliths; the sagittae, lapilli and as-
terisci. Of these, the sagitta generally is the largest, most frequently encoun-
tered, and most relied upon for taxonomic interpretation. Many conclusions
have been drawn regarding the Indians’ fish-eating habits, based upon the
quantity of fish vertebrae (unidentified as to species) present in a given sample
from different levels of the midden. Fitch (1972) states that these conclusions
are untrustworthy, and normally indicate only that fishes were present; they do
not furnish an index of numerical abundance or importance.

A relationship exists between otolith lengths and fish lengths, and once
this relationship has been defined for a given species, a reliable estimate of a
fish size can be ascertained from a single midden otolith. Otoliths can be
deposited in middens by predators through their digestive systems, by scaven-
gers at the midden, or by the Indians. Otoliths that have passed through diges-
tive systems indicate signs of erosion on all surfaces. Species that unlikely
would be captured by the inhabitants for food because of their small size or
living depth could be of predator origin in the midden, and this often can be
determined by the degree of surface erosion. Caution must be used in interpret-
ing these otoliths in relation to the food habits and fisheries of the Indians.
Indians sometimes would consume whole small fishes (Follett 1967), which
offers the possibility that some of the small eroded otoliths could reflect pas-
sage through human digestive systems. Whether these eroded otoliths have
passed through a human or nonhuman digestive system often can be deter-
mined by examining the habits of the fish in question. For example, Fitch
(1969a) noted a myctophid otolith (Ceratoscopelus townsendi ) from a Ventura
midden. The depth at which this lantern fish lives (rarely coming within 600
feet of the surface) as well as its extremely small size (about three inches) was
sufficient evidence to indicate that this fish had not been captured directly by
the inhabitants nor passed through a human digestive system; rather, it was
probably from the stomach of some predator.

METHODS AND MATERIALS

We recovered less than a hundred pounds of midden material from the
site. Our methods of preparing and examining the field sample followed closely
methods discussed by Fitch (1967, 1969a, 1972). The field sample was soaked

1978

Chumash Midden Fish Remains

3

in a large tub of water, then passed through a 14-inch mesh screen. The retained
screenings were set aside to dry. Material not retained by the first screen was
then passed through an 18-mesh screen. These screenings were dried in an
oven at a temperature of 250-300°F. The material that was not retained by this
second screen was passed through a 30-mesh screen. Due to the abundance of
ash and charcoal in the midden “dirt” and its tendency to cling to all other
materials, a secondary screening was necessary. The freshly dried screenings
were soaked separately in a solution of one part liquid bleach and two parts
water for several minutes, then rescreened. This made the screenings cleaner
and reduced them in weight by ten percent. These procedures greatly enhanced
our ability to recognize the fish remains and decreased the time required to
examine the sample. The 14-inch screenings were examined by eye for the
larger fish remains, whereas the 18- and 30-mesh screenings were sorted a
spoonful at a time under a binocular microscope.

HISTORY AND PREVIOUS DIGGING

The Rincon site (SBa-1) became a historic point of contact with the land-
ing of Juan Rodriquez Cabrillo in October 1542. The village, which was called
Xucu by the Indians, was visited several times in succeeding centuries: Sebas-
tian Vizcaino, 1602-1603(7); Gasparde Portola, 1770; and Juan Bautista Anza,
1776. The Rincon area had been ravaged heavily prior to our study. A massive
amount of material was removed by Stephen Bowers, an amateur ar-
chaeologist, in the early 1870’s, but there are no reports on his findings, and the
material subsequently has become lost (Rogers 1929). Leon De Cessac, a
French archaeologist, probably worked ,at the site during the early 1870’s but
these reports remain unverified. Rincon was worked partly in the 1920’s by
archaeologists D.B. Rogers, the Catlin brothers and W.C. Toby and Jesse
Wood. In 1930, Ronald Olson excavated three cemeteries (Olson 1930). Mar-
shall McKusick surveyed the Rincon Point area in 1959 for the University of
California at Los Angeles (UCLA) Archaeological Survey. W.H. Harrison
and Patricia Lyon, then with the University of California, Santa Barbara,
excavated several burials in 1959 and 1960. Patrick Finnerty, formerly of
UCLA, excavated two areas in 1961 and 1964 (Finnerty 1961, 1964). Personnel
representing the UCLA Archaeological Survey have done the remainder of the
excavations: three test pits in April 1963 and extensive excavations in the
summer of 1966. Unfortunately, the control column from the 1966 excavation
was discarded without having been examined. John E. Fitch, California De-
partment of Fish and Game, removed several hundred pounds of material
during the 1966 UCLA excavations and reported three species of fishes: north-
ern anchovy (Engraulis mordax), pacific sardine (Sardinops caeruleus), and
Pacific hake (Merluccius productus). Six other species were found but not
specifically identified (Fitch 1969b). Fitch gave us his material and we have
incorporated it into this report. Most of the Rincon publications report on
aspects other than faunal remains. Evans, Grossman and Toney (1968:36)

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No. 289

mention fish remains but state only that “The remains of fish, mainly verte-
brae, were numerous and appear to represent shallow water species.”

SPECIES ACCOUNTS

It is difficult to present the species accounts in both an archaeological and
an ichthyological approach. We have tried to simplify the matter by separating
the elasmobranchs (Sharks, skates and rays) from the teleosts (bony fishes). In
each of the two groups, taxa are arranged alphabetically by family, and within
each family by genus and species. Common names are listed after each scien-
tific name.

Elasmobranchs
Alopiidae - Thresher sharks

Alopias vulpinus - Thresher shark. — Along the west coast of North
America the thresher shark ranges from central Baja California north to the
Strait of Juan de Fuca, but also is found in all warm seas. Threshers are known
to attain a length of at least 5.5 m (18 feet, Joseph 1954). This shark lives near
the surface, probably not descending to depths greater than 200 m; they rarely
are found near shore. Indians probably utilized a harpoon in obtaining them.
Thresher shark remains have been reported from a midden site in Alameda
County (Follett 1975a).

Material: 1 jaw tooth; Fig. IB.

Carcharhinidae - Requiem sharks

Galeorhinus zyopterus - Soupfin shark. — Soupfin sharks are large, attain-
ing lengths to 2 m (6.5 feet. Miller and Lea 1972). They are found off Chile and
Peru in the southern Hemisphere and from San Juanico Bay, Baja California,
to northern British Columbia in the Northern Hemisphere. South of Point
Conception this shark often inhabits waters several hundred feet deep (Fitch
1968).

Soupfin shark remains also have been recovered from Ventura (Fitch
1969a, 1975; Follett 1965) and Los Angeles counties (Follett 1963a; Frey 1974;
Tartaglia 1976), and at Point St. George site (Gould 1966).

Material: 13 jaw teeth; Fig. II.

Mustelus californicus - Gray smoothhound. — Gray smoothhounds reach
lengths of just over 1.5 m (5 feet, Fitch 1972), and range from Mazatlan,
Mexico, to Cape Mendocino, California. They are common in shallow water
off southern California where they could have been taken easily by gill nets,
hook and line or beach seines, by the Indians.

Remains of gray smoothhounds have been reported from middens in Ven-
tura (Fitch 1969a), Orange (Fitch 1975) and San Diego Counties (Follett 1976).
Material: 2 jaw teeth; Fig. 2A.

Prionace glauca - Blue shark. — The blue shark ranges throughout warm

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Chumash Midden Fish Remains

5

seas. In the eastern Pacific, they are found from Chile to the Gulf of Alaska but
are absent in the tropics. Individuals off California are usually shorter than 1.8
m (6 feet, Fitch 1972). The Indians probably used harpoon and hook and line to
capture this species. Blue shark remains have been recovered also from mid-
dens in Los Angeles (Follett 1963b), San Luis Obispo (Fitch 1972), and Orange
Counties (Fitch 1975).

Material: 1 jaw tooth. Fig. 1H.

Triakis semifasciata - Leopard shark. — The leopard shark has been
taken from Mazatlan, Mexico, to Oregon and in the Gulf of California. They
are common along beaches in bays and attain lengths of 2 m (6.5 feet, Miller
and Lea 1972). The Indians could have captured this shark by harpoon, spear,
hook and line, traps or beach seines.

Leopard shark remains have been recovered from archaeological sites in
Ventura (Fitch 1969a; Follet 1933, 1965), Los Angeles (Follett 1963a, 1963b,
1969), Orange (Fitch 1967, 1975; Follett 1966), Marin, Monterey and San Luis
Obispo (Follett 1964, 1974, 1972a; Fitch 1972), Alameda and Contra Costa
Counties (Follett 1975a, 1975b), and also from Santa Catalina Island (Tartaglia
1976).

Material: 3 jaw teeth, Fig. 2D.

Dasyatidae - Stingrays

Dasyatis dipterura - Diamond stringray. — This large ray ranges from
Paita, Peru, to Kyuquot, British Columbia. They are known to attain a Weight
of 25 kg. (1 13-14 pounds. Miller and Lea 1972). Diamond stingrays are found in
shallow areas to depths of 18 m (55 feet, Miller and Lea 1972) preferring regions
of sandy bottom, where they lie on the substrate and cover themselves with
sand. The Indians could have captured this species with hook and line, spear or
harpoon while fishing in shallow nearshore areas.

The diamond stingray has been reported from only one other midden
(Fitch 1975).

Material: 4 jaw teeth. Fig. 2E.

Heterodontidae - Horn sharks

Heterodontus francisci - Horn shark. — This small nonagressive shark is
found in the Gulf of California and along the coasts of the Californias to
Monterey Bay, reaching a length of just over one meter (3 feet 2-Vs inches,
John Fitch, personal communication). Horn sharks are 'most frequently found
in shallow water in rocky bottom habitats, but they may descend to depths of
164 m (492 feet. Miller and Lea 1972). Fitch (1969a) suggests that horn sharks
may have been captured by the Indians while free diving as well as with hook
and line and possibly with traps.

Horn shark remains have been recovered from Indian middens in Orange
and Ventura Counties (Fitch 1967, 1969a; Follett 1933).

Material: 4 jaw teeth, 1 dorsal spine. Fig. 2F,G,H.

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Hexanchidae - Cow sharks

Notorynchus maculatus - Sevengill shark. — Sevengill sharks range in the
north Pacific from San Carlos Bay, Baja California, to northern British Co-
lumbia. The maximum known length is 2.6 m (8 feet-6 inches, Bohnam 1942).
These sharks commonly are found in bays, but south of Point Conception they
usually inhabit deeper waters of 200 m (600 feet, Fitch 1969a) or more. The
Indians probably captured this species using hook and line in deep water.

Remains of sevengill sharks have been found in middens near Ventura
(Fitch 1969a, 1975) and in Marin County (Follett 1968).

Material: 1 jaw tooth, Fig. IE.

Lamnidae - Mackerel sharks

Carcharodon carcharias - White shark. — The white shark has a
worldwide distribution, and is found in the eastern Pacific from Chile to
Alaska. There is an endemic population off California. They probably are not
uncommon as reported in the literature (Bigelow and Schroeder 1948; Miller
and Lea 1972). Whites are one of the largest sharks attaining lengths of at least
6.4 m (21 feet, Randall 1973). The Indians could have captured this shark by
use of a harpoon.

White shark remains have been reported from sites in Ventura (Fitch 1975;
Follett 1933, 1965) and Los Angeles Counties (Tartaglia 1976), and two sites in
southern New England (Waters 1967).

Material: 3 jaw teeth, Fig. 1 D.

Is urns oxyrinchus - Shortfin mako. — The shortfin mako reportedly at-
tains a length of 3.9 m (13 feet. Miller and Lea 1972) and weight of 454 kg (1000
pounds, Miller and Lea 1972), but a specimen measuring 3.5 m (11 feet-5
inches, Huddleston unpublished data), harpooned off Anacapa Island in Au-
gust and weighing 466 Kg (1030 pounds, Huddleston unpublished data), ap-
pears to be the largest eastern Pacific one (S.P. Applegate, personal communi-
cation). In the eastern Pacific the shortfin mako ranges from Chile to the
Columbia River. The Indians probably caught this shark with harpoon.

Remains of shortfin makos have been recovered previously from midden
sites in Ventura (Fitch 1969a, 1975; Follett 1933, 1965; Tartaglia 1976) and Los
Angeles Counties (Follett 1963a, 1963b, 1969; Tartaglia 1976) and from Santa
Catalina Island (Tartaglia 1976).

Material: 4 jaw teeth. Fig. 1A.

Myliobatidae - Eagle rays

Myliobatis calif ornica - Bat ray. — Bat rays range from the Gulf of
California to Oregon. They have been known to attain a width of about 1 .4 m (4
feet-9 inches, Miller and Lea 1972), and a weight of 95 kg (210 pounds, Miller
and Lea 1972), but most individuals weigh less than 23 kg (50 pounds, Fitch
1969a). They have been found from the surface to depths of 45 m (150 feet.

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Chumash Midden Fish Remains

7

Miller and Lea 1972). Bat rays commonly are found in bays and shallow water
over sandy or muddy bottoms. Large individuals could have been captured by
hook and line, and it is possible that the Indians utilized the caudal stings of
these and other rays for harpoon points.

Bat ray remains have been recovered from several other middens in Ven-
tura (Fitch 1969a, 1975; Follett 1933, 1965; Tartaglia 1976), Los Angeles (Fol-
lett 1963a, 1969; Frey 1974; Meighan 1959; Tartaglia 1976) Orange (Follett
(1966), Marin (Follett 1968, 1974), Santa Barbara (Harrington 1928), San Luis
Obispo Counties (Fitch 1972).

Material: 57 jaw teeth. Figs. 2J,K.

Rhinobatidae - Guitar fishes

Rhinobatos productus - Shovelnose guitarfish. — The shovelnose guitar-
fish is found from the Gulf of California to San Francisco Bay, but recent
reports indicate that it ranges only as far north as Capitola. This cartilaginous
fish has been reported to attain a length of about 1.5 m (61 .5 inches, Miller and
Lea 1972). They are found from the surface to depths of about 13.5 m (45 feet,
Miller and Lea 1972), preferring sandy or muddy bottoms. Sometimes shovel-
nose guitarfish feed in such shallow water as to be nearly beached by the
breaking surf. At such times they are taken easily by spear or hand. It is also
possible that the Indians used hook and line.

Remains of the shovelnose guitarfish have been recovered from other
midden sites in Ventura (Fitch 1969a; Follett 1933; Tartaglia 1976), Orange
(Fitch 1967, 1975), and Los Angeles Counties (Follett 1963a, 1963b; Tartaglia
1976) and Santa Catalina Island (Tartaglia 1976).

Material: 1 jaw tooth. Fig. 2C.

Scyliorhinidae - Cat sharks

Cephaloscyllium ventriosum - Swell sharks. — Swell sharks are found
from Chile to Monterey Bay, including Guadalupe Island and the Gulf of
California. This shark is very abundant around islands south of Monterey Bay,
but not those south of Magdalena Bay. Swell sharks prefer rocky kelp bed
habitat from shallow water to depths of over 400 m (1380 feet, Miller and Lea
1972); they attain lengths of about one meter (3 feet, Miller and Lea 1972).
When caught or threatened they have the ability to inflate their stomachs with
air — a possible defensive mechanism. The Indians probably captured this
shark using traps in shallow rocky areas, although some could have been taken
by hook and line. Swell sharks are reported to be of very poor flavor and the
Indians probably did not actively fish for them (S.P. Applegate, personal
communication).

Swell shark remains also have been recovered from a midden in Ventura
County (Fitch 1969a).

Material: 4 jaw teeth, Fig. 1 F.

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Squalidae - Dogfish sharks

Squalas acanthias - Spiny dogfish. — The spiny dogfish has been found in
the temperate and subarctic regions of both the Atlantic and Pacific Oceans. In
the eastern Pacific, this shark has been found off Chile and from Sebastian
Viscaino Bay, Baja California, to Alaska. They reach lengths of 130 cm (4 feet
3 inches, Bonham, Sanford, Clegg and Bucher 1949), and occasionally travel in
schools in those waters above 390 m (1200 feet, Miller and Lea 1972). Off
California they usually are found in depths between 33 and 66 m (100-250 feet,
Fitch 1969a). The Indians could have utilized hook and line as well as gill nets
to capture this species.

Spiny dogfish remains have been recovered also from Indian middens in
Ventura and San Luis Obispo (Fitch 1969a, 1972), and Los Angeles Counties
(Tartaglia 1976). This species has also been noted from midden sites in British
Columbia, Canada (Niblack 1890; Stwart ms).

Material: 25 jaw teeth. Fig. 1G.

Squatinidae - Angel sharks

Squatina californica - Pacific angel shark. — The Pacific angel shark is
found off Chile and from the Gulf of California to southeastern Alaska, but is
not common north of Point Conception and has not been reported from Canada
(Hart 1973). This shark has been known to reach a length of 1.8 m (5 feet,
Miller and Lea 1972) and a weight of 27 kg (60 pounds, Miller and Lea 1972).
Pacific angel sharks prefer shallow water, dwelling on the bottom in sandy or
muddy areas. They have been observed from depths between about 3 to 45 m
(8 to 150 feet, Fitch 1969a), but prefer a depth of about 20 m. Although the
Indians could have used a beach seine to capture this species they probably
captured them on hook and line or by harpoon.

Remains of the Pacific angel shark have been recovered from middens in
Los Angeles (Follett 1963a, 1963b, 1969; Frey 1974; Tartaglia 1976), Ventura
(Fitch 1969a, 1975; Follett 1965; Tartaglia 1976), Orange (Fitch 1967), and San
Luis Obispo Counties (Fitch 1969a).

Material: 9 jaw teeth, 365 dermal denticles. Fig. 1C.

Unidentified elasmobranch remains

Three fragments of caudal stings were recovered from SBa-1. Unfortu-
nately cuadal stings generally are not useful for making specific identifications
because of their similarity. Two fragments of large caudal stings were recov-
ered from the road level sample near the area where most of the Myliobatis
californica teeth were recovered (Fig. 21, one only). However, they were not
associated with this material and it is possible that they belong to Dasyatis
dipterura , which has a similar type of caudal sting. The third smallest fragment
was found in one of the hill samples and although it could have been from a
small Myliobatis californica it possibly was from Urolophus halleri, the round

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Chumash Midden Fish Remains

9

stingray. Both species are extremely abundant off southern California in shal-
low sandy regions, but because of its small size and fragmented nature the sting
cannot reliably be assigned to either ray.

Material: 3 caudal stings. Fig. 21.

Teleosts

Atherinidae - Silversides

Atherinops affinis - Topsmelt. — Topsmelt range from Santa Maria Bay,
Baja California, to Vancouver Island, British Columbia, and attain a length to
365 mm (Schultz 1933). Topsmelt inhabit bays and kelp beds living at or near
the surface. They possess small mouths and thus would have been difficult to
take by hook and line. The Indians probably used beach seines to obtain this
species.

Fitch (1969a, 1972) reported otoliths A. affinis from sites in Ventura and
San Luis Obispo Counties.

Material: 7 otoliths. Fig. 3 A.

Atherinopsis californiensis - Jacksmelt. — Jacksmelt are known from
Santa Maria Bay, Baja California, to Yaquina Bay, Oregon. Although reported
to reach a length of 22 inches, the largest measured individual was 448 mm
{\1V2 inches, Miller and Lea 1972). Jacksmelt are common along coasts, bays
and kelp beds. Beach seines probably were used to obtain this species.

An otolith of A. californiensis has been reported from a midden near
Ventura (Fitch 1969a); it also has been reported from Marin (Follett 1974), San
Diego (Follett 1976) and Alameda Counties (Follett 1975a).

Material: 3 otoliths. Not figured.

Batrachoididae - Toadfishes

Porichthys notatus - Plainfin midshipman. — Plainfin midshipmen have
been found in the Gulf of California and range from Gorda Bank, Baja Califor-
nia, to Sitka, Alaska. They are found near the surface and to depths of about
330 m (1000 feet. Miller and Lea 1972). South of Point Conception they usually
are confined to deeper waters (Hubbs and Schultz 1939). However, during
“nesting” season it is possible to capture juveniles as well as adults from under
rocks by hand in the intertidal. Plainfin midshipmen spend daylight hours
burrowed in the bottom sediments, emerging at night to seek food. The single
otolith recovered from SBa-1 was that of a juvenile and showed signs of diges-
tive wear; it probably was from the stomach of a predator captured by the
Indians.

Remains oiP. notatus have been reported from middens in Ventura (Fitch
1969a), San Luis Obispo (Fitch 1972), Los Angeles (Fitch 1975) and Alameda
Counties (Follett 1975a).

Material; 1 otolith. Fig. 3C.

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No. 289

Bothidae - Lefteyed flounders

Paralichthys californicus - California halibut. — California halibut are
found from Magdalena Bay, Baja California, to Quillayute River, British Co-
lumbia, and an isolated population occurs in the Gulf of California. They have
been known to reach a length of 1 .8 m (5 feet, Fitch 1969a), and a weight of 33
kg (72 pounds, Fitch 1969a). They are uncommon north of Morro Bay.
California halibut usually are found in shallow water at depths of less than 36 m
preferring sandy habitat. During the spawning season (from February through
July) they frequent shallower water where they could have been captured
easily by hook and line and beach seines.

Remains of California halibut have been reported from middens in Los
Angeles (Follett 1963b, 1969; Tartaglia 1976), Orange (Follett 1966), and Ven-
tura Counties (Fitch 1969a, 1975; Follett 1933; Tartaglia 1976), and from mid-
den sites along the coast of British Columbia (Boas 1895, 1916; Niblack 1890;
Leechman 1973; Ducker 1955, 1965; Sapir 1915; Garfield 1939, 1966; Stewart
ms).

Material: 1 Premaxillary. Fig. 6F.

Carangidae - Jacks and pompanos

Seriola dorsalis - Yellowtail. — The schooling yellowtail range from Chile
to southern Washington and also are found in the Gulf of California. They
attain a length of 1.8 m (5 feet. Miller and Lea 1972) and a weight of 36 kg (80
pounds, Miller and Lea 1972). Generally they live near the surface but may
descend to 24 m. The Indians probably captured this species by hook and line
in offshore areas.

Yellowtail remains have been reported from middens in Los Angeles (Fol-
lett 1963a, 1963b, 1969; Tartaglia 1976) and Ventura Counties (Tartaglia 1976).
Material: 1 vertebra. Not figured.

Trachurus symmetricus - Jack mackerel. — Jack mackerel, a schooling
species, are known from Magdalena Bay, Baja California, to southeastern
Alaska, and from offshore to several hundred kilometers. They sometimes are
found inshore around rocky headlands. Fitch (1972) reported that a 5.25 pound
(2.4 kg) jack mackerel measured 28.5 inches (724 mm) in total length and
possessed otoliths 11.4 mm in length. Trachurus symmetricus is reported to
reach a length of just under one meter (32 inches, Miller and Lea 1972). They
live in the upper 45 m. The Indians probably captured this species with hook
and line or gill nets.

Jack mackerel remains also have been reported in midden sites in Ventura
(Fitch 1969a), San Diego (Fitch 1969b), San Luis Obispo (Fitch 1972) and Los
Angeles Counties (Tartaglia 1976).

Material: 1 otolith. Not figured.

Clinidae - Kelpfishes

Neoclinus uninotatus - Onespot fringehead. — Onespot fringeheads are
known only from Ensenada, Baja California, to Bodega Bay, a restricted geo-

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Chumash Midden Fish Remains

11

graphic distribution. They are a nearshore species occurring in waters 3 to 27 m
(10-90 feet, Miller and Lea 1972) deep, and reach lengths to 247 mm (9 inches,
Miller and Lea 1972). The eroded condition of the otolith recovered from
SBa-1 indicated that it probably had passed through the digestive system of a
predator captured by the Indians. The otolith is from a specimen too small to
have been captured by the gill nets used for white croakers, although many are
taken by hook and line.

N. uninotatus remains have not been reported from any other midden.
Material: 1 otolith. Fig. 3B.

Clupeidae - Herrings

Sardinops caeruleus - Pacific sardine. — Pacific sardines are schooling
fish with a broad distribution in the eastern Pacific from Guay mas, Mexico, to
Kamchatka. They have been known to reach lengths of about 39.4 cm (approx-
imately 16 inches, Miller and Lea 1972). Scales of S. caeruleus are distinctive
among the clupeids. The transverse slits and perforations in the scale are
distinctive for identification. Scales of S. caeruleus were exceedingly abundant
throughout the samples examined with the exception of those from the slope of
Rincon Hill. This species was probably captured by beach seines or gill nets.
Follett (1965, 1968, 1969, 1972a, 1976) reported remains of S. caeruleus (as S.
sagax) from the Conejo Rock Shelter, Tomales Bay site, Century Ranch site,
Mission La Soledad cemetery and a site at Rancho Carrillo, Ventura, Marin,
Los Angeles, Monterey, and San Diego Counties; Fitch (1972, 1975) noted
their remains at sites in San Luis Obispo and Ventura Counties.

Material: 3 otoliths. Fig. 3D.

Embiotocidae - Surfperches

Amphistichus argenteus - Barred surfperch. — Barred surfperch range
from Playa Maria Bay, Baja California, to Bodega Bay, attaining lengths to 43
cm (17 inches, Fitch 1969a) and a weight of two kg (4.5 pounds, Fitch 1969a).
This species is most abundant in breaking surf and other shallow waters, par-
ticularly sandy bottom areas; they rarely are captured in rocky areas. Barred
surfperch have been captured at the surface to depths of 73 m (240 feet, Miller
and Lea 1972). This species probably was captured by beach seines, although
large individuals could have been taken by hook and line.

Barred surfperch remains also have been recovered from midden sites in
Orange (Fitch 1967), Ventura (Fitch 1969a, 1975; Tartaglia 1976; Follett 1933)
and Los Angeles Counties (Tartaglia 1976).

Material: 1 jaw tooth. Fig. 5 A.

Cymatogaster aggregata - Shiner surfperch. — - The shiner surfperch is a
small fish; most individuals are shorter than 177 mm. They range from San
Quintin Bay, Baja California, to Port Wrangell, Alaska, and have been taken
from the surface to depths of about 136 m (480 feet, Miller and Lea 1972).
Shiner surfperch usually prefer water shallower than 18 m. This species, which

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No. 289

does not exceed 100 g i}A pounds, Fitch 1972) in weight, probably was captured
by the Indians using beach seines.

Fitch (1972), reported shiner surfperch otoliths from a midden site at Di-
ablo Cove, San Luis Obispo County.

Material: 2 otoliths. Fig. 3G.

Damalichthys vacca - Pile perch. — The pile perch has been taken from
Guadalupe Island to Port Wrangell, Alaska. They are recorded from surface
waters to depths of 45 m (140 feet, Miller and Lea 1972). A record-sized
specimen measured 44.2 cm (17.4 inches, Fitch 1969a) and weighed slightly
less than 1 .8 kg (4 pounds, Fitch 1969a). The pile perch is found over sandy and
rocky bottom habitat as well as around kelp beds. The Indians probably cap-
tured pile perch using hook and line and possibly gill nets and beach seines.

Follett (1964) found pile perch remains from the Drakes Bay site and Fitch
(1969a, 1972) reported their remains from Ventura and San Luis Obispo Coun-
ties. They also have been reported from midden sites in Marin (Follett 1967),
Monterey (Follett 1973), Ventura and Los Angeles Counties (Tartaglia 1976).
Material: 2 pharyngeal bones, 58 pharyngeal teeth. Fig. 5B,E.

Hyperprosopon argenteum - Walleye surfperch. — This surfperch ranges
from Point Rosarito, Baja California, to Vancouver Island, British Columbia,
including Guadalupe Island. They have a recorded size to 30.5 cm (12 inches.
Miller and Lea 1972). A 27.3 cm (10-% inch, Fitch 1969a) fish weighed just over
397 g (14 ounces, Fitch 1969a). The walleye surfperch has been known from
surface waters to a depth of 18 m (60 feet, Miller and Lea 1972). They prefer
sandy bottom or flat rocky habitat. This species probably was captured by the
Indians using beach seines.

Walleye surfperch remains have been recovered from middens in Ventura
and San Luis Obispo Counties and Drakes Bay (Fitch 1969a, 1972; Follett
1964).

Material: 2 otoliths. Fig. 31.

Phanerodon furcatus - White seaperch. — White seaperch have been
captured from Point Cabras, Baja California, to Vancouver Island, British
Columbia. They have been known from the surface to depths of 42.4 meters
(140 feet, Miller and Lea 1972). This species travels in loose schools over
sandy bottoms. A record-sized specimen weighed about 369 grams (13 ounces,
Fitch 1969a). The Indians probably captured this species using beach seines
near shore and gill nets in deeper waters.

Previously white seaperch have been recovered from a midden in Ventura
County (Fitch 1969a).

Material: 5 otoliths-. Fig. 3H.

Embiotocids — (genus and species undetermined). — Of the 19 species of
surfperches that are found in marine waters off California, 17 occur off south-
ern California. Of these, one is found only around islands, thus there are 16
possible marine species that could have been captured by the inhabitants of
SBa-1. It is possible that some of the 227 unidentified pharyngeal teeth be-

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Chumash Midden Fish Remains

13

longed to some of the other species not listed above or from freshwater
sources.

Material: 227 pharyngeal teeth. Not figured.

Engraulidae - Anchovies

Engraulis mordax - Northern anchovy. — The northern anchovy is one of
the most abundant fishes off our coast. This schooling fish has been found from
Cape San Lucas, Baja California, to Queen Charlotte Island, British Colum-
bia. They are known to attain a length of 229 mm (9 inches, Miller and Lea
1972) but specimens over 177 mm are rare. During fall and winter, northern
achovies apparently move offshore and return inshore during spring (Baxter
1966). During the day, anchovies remain well below the surface, rising to the
surface at night. They are consumed by nearly all predatory fishes off Califor-
nia as well as by birds and marine mammals. The Indians could have caught E.
mordax by beach seine, although some of the otoliths show signs of digestive
wear and were probably from stomachs of predators captured by the Indians.

Northern anchovy remains have been reported from midden sites in
Orange (Fitch 1967), Ventura (Fitch 1969a, 1975) and Los Angeles Counties
(Fitch 1975; Tartaglia 1976). Fitch (1969b) previously reported otoliths of E.
mordax from SBa-1.

Material: 148 otoliths. Fig. 3E.

Labridae - Wrasses

Oxjulis californica - Senorita. — This small wrasse has been taken from
Cedros Island, Baja California, to Sausalito, California. Ranging from surface
waters to depths of 54.5 m (180 feet, Miller and Lea 1972), senorita prefer
shallow water rocky habitat with kelp vegetation. A 229 mm (9-inch, Fitch
1968) individual weighed 113 g (4 ounces, Fitch 1968). The Indians probably
caught this species with gill nets.

O. californica remains have been reported from midden sites in San Luis
Obispo (Fitch 1972), Ventura and Los Angeles Counties (Tartaglia 1976).
Material: 2 otoliths, 7 premaxillary bones, 26 pharyngeal bones. Figs. 3F; 5A;
6C.

Pimelometopon pulchrum - California sheephead. — California
sheephead range from Cape San Lucas, Baja California, to Monterey, with an
isolated population in the northern Gulf of California. Although abundant in
southern California waters, it is not common north of Point Conception. This
fish is abundant in thick kelp beds and slightly rocky habitat. They are known
to reach a length just under one meter (3 feet, Miller and Lea 1972) and a weight
of 16.5 kg (36.25 pounds, Fitch 1968). The Indians could have captured
California sheephead by hook and line, in traps, or by free diving, especially on
offshore islands.

California sheephead remains also have been recovered from many other
midden sites: Arroyo Grande, Point Mugu, Arroyo Sequit, Century Ranch, La

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Jolla, Santa Catalina Island and San Pedro (Wallace 1962; Follett 1933; Mitch-
ell 1959; Follett 1963a, 1963b; Shumway, Hubbs and Moriarty 1961; Meighan
and Eberhart 1953; Meighan 1959; Frey 1974), and from Orange (Fitch 1967),
Ventura (Fitch 1969a; Tartaglia 1976) and Los Angeles Counties (Tartaglia
1976).

Material: 1 jaw tooth. Fig. 5C.

Merlucciidae - Hakes

Merluccius productus - Pacific hake. — The Pacific hake ranges from
Magdalena Bay, Baja California, to Alaska and along the Asiatic coast; there is
an isolated population in the Gulf of California. They have been known to
reach a length of nearly one meter (3 feet, Fitch 1972) and a weight of 3.6 to 4.5
kg (8 to 10 pounds, Fitch 1972). Although found from the surface to depths of
1000 m (3000 feet, Miller and Lea 1972), most individuals off California inhabit
waters shallower than 270 m (750 feet, Fitch 1972). The Pacific hake is a
schooling fish that moves inshore and offshore much the same as the northern
anchovy, and is preyed upon heavily by marine mammals. No Pacific hake
remains were recovered from the samples examined by us, but Fitch (1969b)
reported jaw fragments and vertebrae of Pacific hake from a sample (general
sample) he examined from the slope region of Rincon Hill.

Pacific hake remains have been recovered from Indian midden sites in
Ventura and San Luis Obispo Counties and at Tomales Bay and Point St.
George (Fitch 1969a, 1969b, 1972; Follett 1968; and Gould 1966).

Material: Jaws, vertebrae. Not figured.

Sciaenidae - Croakers

Cy noscion nobilis - White seabass. — White seabass are found from Mag-
delena Bay, Baja California, to Juneau, Alaska, with an isolated population in
the northern Gulf of California. Although reported to grow as large as 41 kg (90
pounds. Miller and Lea 1972), a 38 kg (83.75 pound, Fitch and Lavenberg 1971)
individual is recognized as the world record. The white seabass prefers depths
of 21 to 45 m (75 to 150 feet, Fitch and Lavenberg 1971) but has been taken
from surface waters to depths of 121 m (400 feet, Miller and Lea 1972). Al-
though large adults have few natural enemies, the largest of the two sagittae
that we recovered at SBa-1 was eroded, indicative of digestive wear possibly
due to consumption by a large predator. Otoliths of C. nobilis have been used
by California’s Indians for necklace ornaments (Fig. 7). The Chumash proba-
bly caught white seabass on hook and line while fishing just offshore.

C. nobilis remains have been found at Malaga Cove and Santa Catalina
Island (Walker 1951; Meighan 1959), Arroyo Sequit and Century Ranch sites
(Follett 1963a, 1963b; Mitchell 1959), Ventura (Fitch 1969a, 1975; Tartaglia
1976), San Nicolas Island (Charles Rozaire, Natural History Museum of Los
Angeles County, personal communication), Marin (Follett 1974), Contra
Costa (Follett 1975b) and Los Angeles Counties (Tartaglia 1976), and a midden

1978

Chumash Midden Fish Remains

15

site at Punta Pequena, San Juanico Bay, Baja California (Huddleston, unpub-
lished data).

Material: 2 otoliths. Figs. 4 A, B.

Genyonemus line at us - White croaker. — White croakers attain lengths to
412 mm (15.4 inches, Miller and Lea 1972). They have been found from Mag-
dalena Bay, Baja California, to Vancouver Island, British Columbia, but are
not common north of San Francisco Bay. White croakers prefer sandy or
muddy bottom habitat and frequently are taken in shallow waters, although
they have been known from depths of 188 m (330 feet. Miller and Lea 1972). A
random sample of 100 G. lineatus otoliths from SBa-1 ranged in size from 7.0 to
12.5 mm representing fish from 152 to 266 mm in length. This narrow size range
indicates a selective method of fishing, probably the use of gill nets.

Remains of G. lineatus were reported previously from SBa-1 and near
Ventura (Fitch 1969b, 1969a, 1975), Conejo Rock shelter and a site near La
Jolla (Follett 1965; Shumway et al. 1961), San Diego County (Follett 1976) and
from a site in San Pedro (Frey 1974).

Material: 932 otoliths. Fig. 4C.

Seriphus politus - Queenfish. — This fish is found from west of Uncle
Sam Bank, Baja California, to Yaquina Bay Oregon, but is rare north of Mon-
terey. Queenfish attain lengths of 305 mm (12 inches Fitch 1972) and a weight
just over 284 g (10 ounces, Fitch 1972). They prefer much the same habitat as
the white croaker, and most of the individuals probably were captured in gill
nets along with the white croakers. Queenfish are not taken as easily by hook
and line as white croakers, but it is possible that some were captured that way.

Fitch (1969a, 1972, 1975) reported S. politus otoliths from sites in Ventura
and San Luis Obispo Counties.

Material: 42 otoliths. Fig. 4D.

Scombridae - Mackerels and Tunas

Pneumatophorus japonicus - Pacific mackerel. — This transpacific
species is found in the eastern Pacific from Chile to the Gulf of Alaska. Attain-
ing a length of 635 mm (25 inches, Miller and Lea 1972) and a weight of 2.9 kg
(6-16 pounds, Miller and Lea 1972), this schooling species is found from the
surface to depths of 45.5 m (150 feet, Miller and Lea 1972). The Pacific mac-
kerel, like the bonito, will take nearly any bait, and the Indians probably
captured them by the use of hook and line.

Follett (1963a, 1963b, 1965) reported Pacific mackerel remains from two
sites in Los Angeles County and from the Conejo Rock Shelter in Ventura
County. Fitch (1969a, 1975) reported their remains from midden sites near
Ventura, and Follett (1976) reported their occurrence from a site in San Diego
County. They have also been reported from a midden site at Cape Brinera
(Sidimi) USSR (Besednov 1973).

Material: 4 otoliths. Figs. 4E.; 6 A.

Sarda chiliensis - Pacific bonito. — - The schooling Pacific bonito ranges

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from Chile to the Gulf of Alaska but is absent in tropical waters and uncommon
north of Point Conception. They have been reported to attain a length of just
over one meter (40 inches. Miller and Lea 1972). A 686-mm (27 inches, Fitch
1969a) specimen weighed 4.8 kg (\O-V2 pounds, Fitch 1969a). This fish usually
travels at or near the surface and will take nearly any bait. The Indians proba-
bly captured Pacific bonito with hook and line. Follett (1933, 1963a, 1963b)
reported Pacific bonito remains from two middens in Los Angeles County and
from one in Ventura County. Fitch (1969a) noted their remains from a Ventura
County midden and Tartaglia (1976) noted them from sites in Ventura and Los
Angeles Counties.

Material: 33 dentaries. Fig. 6B.

Scorpaenidae - Rockfishes

Sebastes atrovirens - Kelp rockfish. — The kelp rockfish ranges from Pt.
San Pablo, Baja California, to Timber Cove, Sonoma County. They have been
known to attain a length of 425 mm (16-3A inches, Fitch 1969a) and a weight of
just over 1.4 kg (3 pounds, Fitch 1969a). Kelp rockfish inhabit depths from
subtidal to 45.5 m (25 fathoms, Phillips 1957) but are most abundant at about 10
m or less. The inhabitants of SBa-1 probably caught kelp rockfish by hook and
line in shallow water just offshore or in traps.

Remains of S. atrovirens also have been reported from a midden in San
Luis Obispo County (Fitch 1972).

Material: 1 otolith. Fig. 4F.

Sebastes diploproa - Splitnose rockfish. — This deep-living rockfish
ranges from north of San Martin Island, Baja California, to Prince William
Sound, Alaska. They are known to reach a length of 457 mm (18 inches, Miller
and Lea 1972) and a depth of 473 m (250 fathoms, Phillips 1957). The single
broken otolith recovered from SBa-1 showed little digestive wear. Because this
species has not been known to come nearer the surface than 212 m (700 feet,
Miller and Lea 1972) it may not have been captured directly by the Indians. We
suspect that it was derived from the digestive system of a deep-feeding pre-
dator such as a marine mammal, which was captured by the Indians.

Remains of S. diploproa have not been reported from any other midden.
Material: 1 otolith. Fig. 4H.

Sebastes goodei - Chilipepper. — S. goodei have been captured from
Magdalena Bay, Baja California, to near Cape Scott on the northwest coast of
Vancouver Island, British Columbia. They range from the surface to great
depths. The deepest recorded specimen was taken at 327 m (1080 feet, Miller
and Lea 1972); the largest specimen captured measured 559 mm (22 inches,
Phillips 1957). Chilipeppers frequently are captured by sport fishermen, but
only from skiffs or boats anchored or drifting over deep rocky habitat. Al-
though young individuals usually remain in shallow water, all of the otoliths we
recovered at SBa-1 were from adult fish. The Indians probably captured this
fish by hook and line while fishing in deep water.

1978

Chumash Midden Fish Remains

17

Follett (1963a) reported S. goodei from a midden in Los Angeles County.
Material: 9 otoliths. Fig. 4G.

Sebastes miniatus - Vermilion rockfish. — Ranging from San Bonito Is-
land, Baja California, to Vancouver Island, British Columbia, this rockfish has
been captured at a depth of 200 m (1 10 fathoms, Phillips 1957); juveniles prefer
shallow water. Vermilion rockfish attain lengths of 762 mm (30 inches, Phillips
1957). The Indians probably caught this species while fishing with hook and
line in intermediate depths. Remains of vermilion rockfish have been recov-
ered from Indian middens at Pt. St. George and Scripps Estate (Gould 1966;
Shumway et al. 1961).

Material: 2 otoliths. Fig. 41.

Sebastes spp. - (Species undetermined). — At least 58 species of
rockfishes (genus Sebastes ) inhabit waters off California. Some of the frag-
mented Sebastes otoliths we recovered from SBa-1 probably belonged to one
or more of these species.

Serranidae - Sea basses

Stereolepis gigas - Giant sea bass. — Sometimes placed in the family
Percichthyidae (temperate basses), the giant sea bass has been taken from the
Gulf of California to Humbolt Bay, but is not abundant north of the Channel
Islands off southern California. A record specimen weighing 252 kg (557
pounds, Fitch and Lavenberg 1971) was caught in 1962. An individual weighing
197 kg (435 pounds, Fitch and Lavenberg 1971) was found to be 72 to 75 years
old (Fitch and Lavenberg 1971). Large individuals prefer rocky bottom habitat
and depths of 35 to 46 m, just outside kelp beds. During the spawning period,
which is from June to September, large individuals are relatively abundant in
shallow waters where they can be taken easily with hook and line.

Fitch (1969a) reported branchiostegal rays of S. gigas from a midden near
Ventura.

Material: 1 vertebra. Fig. 51.

Sphyraenidae - Barracudas

Sphyraena argentea - Pacific barracuda. — Ranging from Cape San
Lucas, Baja California, to Kodiak Island, Alaska, the Pacific barracuda
reaches a length of 1 .2 m (46-Ji inches, Fitch and Lavenberg 1971) and a weight
of 7.7 kg (18 pounds 3 ounces, Fitch and Lavenberg 1971). There is an un-
verified record of 1.5 m. Most common south of Morro Bay, this schooling
species remains near shore in depths from the surface to 18.8 m (60 feet. Miller
and Lea 1972). Schools of juveniles will sometimes wander into shallow bays.
Although today barracudas often are captured by gill nets, the inhabitants of
SBa-1 probably took barracudas by hook and line.

Remains of Pacific barracuda have been recovered from midden sites in
Ventura (Fitch 1969a, 1975; Follett 1933, 1965; Tartaglia 1976), Los Angeles

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No. 289

(Follett 1963b, 1969; Tartaglia 1976) and San Diego Counties (Follett 1976).
Material: 6 otoliths. Figs. 5D, 6C,D,E.

Xiphiidae - Swordfish

Xiphias gladius - Swordfish. — This species is found worldwide in warm
and temperate seas, but in the eastern Pacific it is found from Chile to Oregon.
The swordfish occurs off our coast only during a few summer months each
year. They frequently rest at the surface but have been seen at depths greater
than 666 m (2000 feet, Fitch and Lavenberg 1971). The largest recorded speci-
men measured 4.5 m (14 feet 1 1-14 inches, Fitch and Lavenberg 1971) and
weighed 535 kg (1182 pounds, Fitch and Lavenberg 1971). This specimen was
captured off the coast of Chile in 1953. The Indians probably obtained
swordfish with harpoons during summer months.

Swordfish remains have been reported from midden sites in Ventura
County (Fitch 1969a; Follett 1933) and from the Burton Mound Site (Har-
rington 1928).

Material: 7 vertebrae. Figs. 5F,G.

ELEMENTS OF IDENTIFICATION

Otoliths were by far the most important element in identifying teleosts
from SBa-1. Twenty of the 29 species (representing 69 percent) were identified
from otoliths; the next most useful elements were premaxillaries, pharyngeal
bones, dentaries and vertebrae. Four species were identifiable by each of these
elements, representing only 14 percent of the teleost species. For identifying
elasmobranchs, which do not possess otoliths, teeth were the most important
element. All 16 of the sharks were identifiable on the basis of teeth alone. One
species also was identified from dermal denticles and another by a dorsal fin
spine, each representing less than 6.5 percent of the elasmobranch species.
Small mesh screens are not only necessary for retaining small otoliths, but are
important in the collecting of elasmobranch remains. From the lA- inch
screenings a total of four species could be identified; from the 18-mesh, eight
species; and from the 30-mesh, 11 species. Six species of four families were
retained only by the 30-mesh screen. Utilizing screens smaller than 30- or
32-mesh is not practical. There are otoliths which will pass through such
screens, but they represent fishes that would not likely have been utilized as
food by the Indians who inhabited these sites. It would be desirable to examine
a large midden sample with 30-mesh screens, but if a shortage of time makes
this impossible at least a 6 x 6-inch control column should be given this fine
screening examination.

OTOLITHS IN ORNAMENTATION

A necklace in the Natural History Museum of Los Angeles County
(LACM) no. A. 5600/99 from a San Nicolas Island site, mentioned by Fitch

1978

Chumash Midden Fish Remains

19

Scientific name

TABLE 1

a

<u

x oo
2 e
S be
2 j= o

CX CX-O

e

>

C3 0)

2.S

o cx

Elasmobranchs

Alopias vulpinus
Carcharodon carcharias
Cephaloscyllium ventriosum
Dasyatis dipterura
Galeorhinus zyopterus
Heterodontus francisci
Isurus oxyrinchus
Mustelus californicus
Myliobatis californica
Myliobatoidei
Notorynchus maculatus
Prionace glauca
Raja spp.

Rhinobatos productus
Saualus acanthias
Squatina californica

Teleosts

Amphistichus argenteus
Atherinops affinis 7

Atherinopsis californiensis 3

Cymatogaster aggregata 2

Cynoscion nobilis 2

Damalichthys vacca
Engraulis mordax 148

Embiotocids

Genyonemus lineatus 932

Hyperprosopon argenteum 2

Merluccius productus
Neoclinus uninotatus 1

Oxyjulis californica 2

Paralichthys californicus
Phanerodon furcatus 5

Pimelometopon pulchrum
Pneumatophorus japonicus 4

Porichthys notatus 1

Sarda chiliensis

Sardinops caeruleus 3

Sebastes atrovirens 1

Sebastes diploproa 1

Sebastes goodei 9

Sebastes miniatus 2

Sebastes spp. X

1

3

4
4

13

4

4

2

57

1

1

12 1*

1

25

9 395**

58

2

X

7 26

1

9

33

X

2

20

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No. 289

TABLE 1 (Continued)

Scientific name

o

Teleosts (continued)

Seriola dorsalis
Seriphus politus

42

Sphyraena argentea
Trachurus symmetricus
Xiphias gladius

6 X

X X

7

X

(*) Wing spines.

(**) Dermal denticles.

(1969a), is constructed of white seabass otoliths and Olivella shells (Fig. 7).
The 30 otoliths (14 right sagittae + 16 left sagittae) have been drilled through
the thick knob-like protrusion on the outer face. The otoliths apparently were
drilled part way through on one side and then turned over and completed from
the other side. The hardness of otoliths as well as their vulnerability to fracture
when drilling pressure is applied indicates that great patience was required to
complete this necklace. None of the otoliths from SBa-1 showed any signs of
such drilling or unnatural modification.

Also in the LACM collection is an artifact (no. L.2100. A. 902.70-1) from
another midden in Los Angeles County (LAn-174) in which white croaker
otoliths were used in a decoration (Fig. 8) believed to be a portion of a water
jug (C. Rozaire, personal communication). Small pebbles and shell fragments
are embedded randomly in the asphalt; these are probably a natural encase-
ment. A faint fabric-like pattern is detectable on part of the asphalt surface.
Sixteen white croaker sagittae (22 present in an earlier photograph) also are
embedded in the asphalt in an “L” -shaped pattern. All of the otoliths are
embedded with the outer face showing and nearly all with the end pointing
inward. There is no apparent pattern in the placement of right and left sagittae.
Three white croaker sagittae were loose in the bottom of the tray and several
“empty” otolith impressions can be seen in the “L” shaped pattern in the
asphalt. It is assumed that the otoliths are merely a form of decoration; no
other explanation is suggested at present. None of the otoliths recovered from
SBa-1 shows any asphalt or tar residues to suggest their possible use as orna-
ments, trinkets or fetishes.

COMPARISON BETWEEN THE FISH REMAINS AND FISHING
TECHNIQUES OF SBa-1 AND OTHER SITES

The oceanic environment adjacent to SBa-1 and Ven-3 in Ventura County
is nearly identical, both sites possessing offshore kelp beds, sandy regions and

1978

Chumash Midden Fish Remains

21

nearby rocky habitat; and similar species of fishes inhabit their coastal regions.
The similarity of the fishing cultures of these two villages is reflected in the fish
remains (Table 2). Thirty-one (72 percent) of the 41 species from SBa-1 (which
likely were captured by the inhabitants as food items) also were recovered from
the Ven-3 midden. Of the species thought to have been captured directly by the
inhabitants, 48 percent from SBa-1 probably were captured by hook and line
compared to 52 percent from Ven-3. Twenty-eight percent of the species of
SBa-1 probably were captured by gill nets compared to 33 percent from Ven-3.
Probable beach seine captures accounted for 40 percent of the species from
SBa-1 and 44 percent of the species from Ven-3. Eight percent of the species of
SBa-1 were captured by other methods compared with 11 percent of the
species of Ven-3. These figures are only approximate since in several cases
some of the species probably were captured by more than one method. Unfor-
tunately, there are no age-data correlations between our samples from SBa-1
and those from the Ven-3 site reported by Fitch (1969a). It is apparent that
these two villages were utilizing similar fishing techniques and were oriented
strongly towards a marine fishery. This is in contrast with the inhabitants of
Ora- 190 in Orange County, who apparently utilized only hook and line and
traps in taking fish (Fitch 1967). The inhabitants of SLO-2, San Luis Obispo
County, also were apparently not fishery oriented, although they occupied the
same site for several thousand years: “There is no evidence that they prog-
ressed beyond a hook and line fishery . . (Fitch 1972).

ELASMOBRANCH REMAINS

The relative scarcity of shark teeth in the SBa-1 midden when compared to
the total number of vertebrae recovered is perhaps the result of one or two
factors. First, sharks could have been decapitated at or near their point of
capture or at a “cleaning station”, reducing the total weight to be carried and
the chance of being “bitten”. Second, the teeth could have been utilized in
some form of ornamentation or other artifact usage, but there is no indication
of this.

Analysis of Fish Remains

In attempting to analyze the fishery habits of the inhabitants of SBa-1 we
selected only the bony fishes for which an approximate minimum number of
individual (MNI) fish could be determined. Further, we utilized only those
species that were presumed to be food sources. We omitted the anchovies
because it was not possible to determine how many of the otoliths were from
stomachs of predators captured by the Indians. The number of otoliths for each
species was divided by two (since each fish has two sagittae), a practice which
assumes that both right and left sagittae were recovered, but if not true, estab-
lishes a minimum number. For other elements (identifications based solely on
vertebrae, etc.) only a MNI of one was interpreted, although in the case of the

22

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No. 289

TABLE 2

Probable method of capture

Hook

Scientific name Occurrence and Beach Gill

in Ven-3 line seine net Other

Elasmobranchs

Alopias vulpinus
Carcharodon carcharias
Cephaloscyllium ventriosum
Dasyatis dipterura
Galeorhinus zyopterus
Heterodontus francisci
Isurus oxyrinchus
Mustelus californicus
Myliobatis californica
Notorynchus maculatus
Prionace glauca
Raja spp.

Rhinobatos productus
Squalus acanthias
Squatina californica
Triakis semifasciata

Teleosts

Amphistichus argenteus
Atherinopos affinis
Atherinopsis californiensis
Cymatogaster aggregata
Cynoscion nobilis
Damalichthys vacca
Engraulis mordax
Genyonemus lineatus
Hyperprosopon argenteum
Merluccius productus
Oxyjulis californica
Paralichthys californicus
Phanerodon furcatus
Pimelometopon pulchrum
Pneumatophorus japonicus
Sarda chiliensis
Sardinops caeruleus
Sebastes atrovirens
Sebastes goodei
Sebastes miniatus
Seriphus politus
Sphyraena argentea
Stereolepis gigas
Trachurus symmetricus
Xiphias gladius

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X X

X

X X

X

X

X

1978

Chumash Midden Fish Remains

23

swordfish the seven vertebrae could have come from seven different individuals.

Since it was not possible to determine the MNI for the elasmobranchs
because of the abundance of teeth in a single jaw, no attempt was made to
indicate preference or selective fishing for this group. According to present-
day taste standards the thresher shark is considered choice, followed closely
by the white shark and the shortfm mako, but all of these are fast-swimming
forms and not easily taken (S.P. Applegate, personal communication).

A minimum of 538 bony fishes was represented by the teleost remains.
Although we considered 23 species to be potential food fishes, 19 of these
constituted only 6.4 percent MNI; whereas white croaker constituted 86.6
percent MNI. Single representatives of such large fishes as the giant sea bass
and swordfish would yield large poundage of food, but the relative scarcity of
their remains in SBa-1 tends to indicate only limited or occasional usage and we
do not believe that they were relied upon heavily for food. The large percent-
age of white croaker suggests strongly that the Indians either preferred this
species or else it was easy to catch and not undesirable.

Although a hook and line fishery would account for the greatest diversity
of teleostean species when compared with other fishing methods used by the
Indians, it could not have accounted for more than 6.6 percent of the MNI.
Beach seining probably took the second greatest number of species, but only
2.76 percent MNI. Fewer species were taken by gill netting but this method
apparently accounted for nearly 92 percent MNI. All other methods yielded
only a fraction of one percent MNI. Gill netting is indicated as the primary
method of fish gathering.

DISCUSSION

Fish remains were very scarce in all samples obtained from the slope of
Rincon Hill. The slope area is believed to be an older region of occupation,
estimated to be 2000 to 1000 B.C. (Evans et al. 1968). Three radiocarbon dates
obtained from samples collected by Lyon and Harrison in 1959-60, were 1320
B.C. ± 250 years, 1470 B.C. ± 130 years and 1580 B.C. ± 60 years (Radiocar-
bon 1963:290). Sufficient fish remains were recovered from Rincon Hill to
indicate that fish were utilized in the Indians’ diet. The lack of variety, the
absence of open ocean or deep dwelling species, and the overall sparseness of
the fish remains indicate a minimal usage of fish as a food source. This proba-
bly reflects inadequate technical skills or inefficient fishing methods at that
stage of cultural development.

This is in contrast with the lower and more recently occupied region of the
SBa-1 complex. Based on the fish material we recovered the occupants of the
lower area displayed highly advanced fishing techniques and depended upon
the sea as a primary source of food. The occupation of this lower region
pen6isted into historic times.

Among the samples from the lower region were remains of many pelagic or
open ocean fishes, species which could not have been captured from shore.

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No. 289

Contributions in Science

Based on the life habits of the fishes (presumably these habits have not changed
in the last several thousand years), the inhabitants of this lower region were
utilizing crafts to carry them well beyond the surf zone and kelp beds where
they harpooned swordfish, white sharks, thresher sharks, blue sharks and
shortfin makos. With deep hook and line fishing they were able to take skates,
spiny dogfish and rockfish; closer to shore, hook and line methods were practi-
cal in kelp bed areas to obtain California sheephead. In shallow waters and surf
zones beach seines would explain the presence of smaller-mouthed fishes such
as topsmelt, jacksmelt, and some of the small surfperches as well as bat rays
and sting rays. Use of gill nets was demonstrated by the abundance of white
croaker otoliths of nearly identical sizes. Fishes that inhabit shallow water and
are approached easily, such as the shovelnose guitarfish and the horn shark,
could have been taken by hand or spear.

A variety of fishing gear has been reported for the Chumash: plank
canoes, harpoons, fish spears, harpoon arrows, shellfish hooks, bone fish
hooks, traps and nets (Hoover 1973; Richie and Hager 1973). Concerning the
SBa-1 sites only shell fish hooks have been reported (Evans et al. 1968). Evans
et al. (1968:24-25) described a 1 ‘donut-shaped stone” as follows: “One ar-
tifact ... is a round cobble which has had a cylindrical hole (which) contains a
large amount of asphaltum.” This stone was nine centimeters in diameter, 6.8
cm thick and had a bore diameter of 3 cm. It is obvious from this description
that they were dealing with a stone fishing weight (sinker) probably similar to
those illustrated by Hoover (1973, plate 4, fig. B and C). Vague descriptions of
“pointed bone artifacts” and “hammered stones” by Evans et al. (1968) ren-
der it impossible to determine if some of these materials were remnants or
represented bone fish hooks, bone gorges, or fishing weights. Such artifacts
should have been present at the site.

Olson (1971) noted that “ear bones” could at times be of value in age
determination but failed to mention their significance in establishing specific
identifications. Further, he suggests that the lack of fish remains in some sites
could be due to the possible cartilaginous skeletal elements such as found in
Salmo; this theory overlooks the fact that Salmo otoliths, as with most otoliths,
are aragonitic in composition. Even fishes which possess poorly ossified
skeletal structures, if present in the midden, would leave behind their otoliths.

In his discussion on shark remains, Olsen (1971:2-6) states, “A number of
cartilaginous fish are poisonous when eaten by man, and it is not known
whether the recovered shark and ray elements are the residue of meals or
represent some other use by man.” S.P. Applegate (personal communication)
states that none of the sharks or rays occurring in waters off North America is
poisonous when eaten by man. Therefore, there is no reason to assume that the
shark and rays recognized in a midden site do not constitute food items.

Unfortunately the randomness of our samples and the disrupted state of
the Rincon site (SBa-1) prevented a study of an interesting transgression of the
SBa-1 inhabitants from a terrestrial hunting to a marine-oriented people.

1978

Chumash Midden Fish Remains

25

Figure 1. A lsurus oxyrinchus tooth, 18.9 mm high; B. Alopias vulpinus tooth, 6.0 mm
high; C. Squatina californica tooth, 3.8 mm high; D. Carcharodon carcharias tooth, 22.5
mm high; E. Notorynchus maculatus incomplete tooth, 9.0 mm length of base; F.
Cephaloscyllium ventriosum tooth, 2.7 mm high; G. Squalus acanthias tooth, 3.7 mm
length of base; H. Prionace glauca worn tooth, 8.5 mm high; I. Galeorhinus zyopterus
tooth, 4.1 mm high.

Contributions in Science

No. 289

Figure 2. A. Mustelus californicus tooth, 1.1 mm high; B. Raja sp. tooth, 2.1 mm high;
C. Rhinobatos productus tooth, 1.4 mm high; D. Triakis semifasciata tooth, 3.2 mm
high; E. Dasyatis dipterura tooth, 1.7 mm high; F . Heterodontus francisci anterior tooth,
2.9 mm high; G. Heterodontus francisci posterior tooth, 7.6 mm high; H. Heterodontus
francisci incomplete dorsal spine, 31.4 mm; I. Myliobatoidei incomplete caudal sting,
49.0 mm; J. Myliobatis californica dorsal view median tooth, 26.3 mm length of base; K.
Myliobatis californica ventral view median tooth.

1978

Chumash Midden Fish Remains

27

Figure 3. A. Atherinops affinis left sagitta, 5.1 mm; B. Neoclinus uninotatus left sagitta,
1.9 mm; C. Porichthys notatus eroded right sagitta, 1.3 mm; D. Sardinops caeruleus
right sagitta, 3.5 mm; E. Engraulis mordax right sagitta, 4.0 mm; F. Oxyjulis californica
left sagitta, 1.9 mm; G. Cymatogaster aggregata right sagitta, 6.7 mm; H. Phanerodon
furcatus right sagitta, 8.4 mm; I. Hyperprosopon argenteum right sagitta, 6.7 mm.

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Contributions in Science

No. 289

Figure 4. A. Cynoscion nobilis eroded left sagitta, 25.7 mm; B. Cynoscion nobilis left
sagitta, 21.1 mm; C. Seriphus politus left sagitta, 9.3 mm; D. Genyonemus lineatus left
sagitta, 18.0 mm; E. Pneumatophorus japonicus left sagitta, 5.3 mm; F. Sebastes atrovi-
rens right sagitta, 10.1 mm; G. Sebastes goodei left sagitta, 19.8 mm; H. Sebastes
diploproa eroded right sagitta rostrum missing, 10.2 mm; I. Sebastes miniatus right
sagitta, 18.7 mm.

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Chumash Midden Fish Remains

29

Figure 5. A. A mphistichus argenteus pharyngeal tooth, 4.8 mm; B. Damalichthys vacca
pharyngeal tooth, 6.0 mm high; C. Piomelometopon pulchrum tooth, 7.3 mm high; D.
Sphyraena argentea left sagitta, 15.2 mm; E. Damalichthys vacca lower pharyngeal bone,
26.2 mm wide; F. Xiphias gladius vertebra, 46.9 mm length; G. Xiphias gladius end view
of vertebra, 47.6 mm diameter; H. Oxyjulis californica lower pharyngeal bone, 5.2 mm
wide; I. Stereolepis gigas vertebra, 60.0 mm high.

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Contributions in Science

No. 289

Figure 6. A. Pneumatophorus japonicus incomplete right dentary, 42.9 mm B. Sarda
chiliensis left dentary, 56.3 mm; C. Sphyraena argentea right dentary, 64.4 mm; D.
Sphyraena argentea right palatine, 51.8 mm; E. Sphyraena argentea incomplete right
pre-maxillary, 30.3 mm; F. Paralichthys californicus incomplete left pre-maxillary, 45.1
mm; G. Oxyjulis californica right dentary, 10.9 mm.

1978

Chumash Midden Fish Remains

31

Figure 7. Necklace, LACM no. 5600/99, constructed with Cynoscion nobilis otoliths
and Olivella shells.

32

Contributions in Science

No. 289

Figure 8. Fragment of “water jug” LACM no. L. 2100. A. 902. 70-1 containing otoliths of
Genyonemus line at us.

1978

Chumash Midden Fish Remains

33

ACKNOWLEDGMENTS

We wish to thank the following individuals for their assistance and contri-
butions in our studies: Shelton P. Applegate, for verification of the elasmo-
branch remains; R.O. Browne, C.H.Eugler, Frances A. Fitch and Roberta
Greenwood who were important for John Fitch’s samples; Leonard Nelson,
University of California, Los Angeles, for historical information on the SBa-1
site; Mark Roeder, Natural History Museum of Los Angeles County, and
Bruce Welton, University of California, Berkeley, for their aid in the elasmo-
branch identifications: Camm C. Swift, Natural History Museum of Los
Angeles County, for his aid in the identification of the teleost material; Robert
J. Lavenberg, Natural History Museum of Los Angeles County, for his helpful
comments in the construction of this manuscript.

We especially wish to thank John E. Fitch, California Department of Fish
and Game, for incorporation of his site samples into our study, for confirma-
tion of otolith identifications, and for his encouragement and invaluable advice
throughout our study.

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Besednov, L.N. 1973. Food fish of Sidimi Bay (an inlet of Peter the Great Bay) in the
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Evans, Robert K., Joel Grossman and James T. Toney. 1968. Archaeological Salvage
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Fitch, John E. 1967. Fish remains recovered from a Corona del Mar, California, Indian
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at Diablo Cove, San Luis Obispo County, California. San Luis Obispo Co., Ar-
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. 1975. Fish remains from a Chumash village site (Ven-87) at Ventura, Califor-
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Fitch, John E., and Robert J. Lavenberg. 1971. Marine food and game fishes of
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Follett, W.I. 1933. Incomplete list of fishes from Point Mugu shellmound. Unpublished
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. 1957. Fish remains from a shellmound in Marin County, California. American

Antiquity, Vol. 23, No. 1:68-71.

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County, California, Calif. Dept. Parks and Recreation, Div. Beaches and Parks,
Archeol. Dept., 9:113-121.

. 1963b. Fish remains from the Century Ranch site (LAn-227), Los Angeles

County, California. Univ. Calif. Los Angeles, Archeol. Surv., Rept. 1962-
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. 1964. Fish remains from a sixteenth century site on Drakes Bay, California.

Univ. Calif. Los Angeles, Archeol. Surv., Ann. Rept. 1963-1964:27-41.

. 1965. Fish remains from the Conejo Rock Shelter site (Ven-69), Ventura

County, California. Univ. Calif. Los Angeles, Archeol. Surv. Ann. Rept., 1964-
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. 1966. Fish remains from archaeological sites at Irvine, Orange County,

California. Univ. Calif. Los Angeles, Archeol. Surv. Rept., 1965-1966:185-195.

. 1967. Fish remains from coprolites and midden deposits at Lovelock Cave,

Churchill County, Nevada. Repts. Univ. Calif. Archeol. Surv., Berkeley, 70:93-
116.

. 1968. Fish remains from two submerged deposits in Tomales Bay, Marin

County, California. Occas. Pap., Calif. Acad. Sci., 67:1-8.

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California. Univ. Calif. Los Angeles, Archeol. Surv. Ann. Rept., 1968, 10:32-143.

. 1972a. Fish remains from Mission La Soledad Cemetry, Mnt-223, Monterey

County, California. Monterey Co. Archeol. Soc. Quart., Vol. 1(3): 11.

. 1972b. Fish remains from the Kodani Site (Mnt-436, Monterey County,

California). Monterey Co. Archeol. Soc. Quart, Vol. I(4):3-4.

1978

Chumash Midden Fish Remains

35

. 1973. Fish remains of the Church Rockshelter Mnt-44, Monterey County,

California. Monterey Co. Archeol. Soc. Quart, Vol. 11(4): 10-11.

. 1974. Fish remains from the Shelter Hill Site, Marin County, California.

Appendix X. San Francisco State University Treganza Anthro. Mus. Pap. 15
(MAPOM pap.2): 145-159.

. 1975a. Fish remains from the west Berkeley shellmound (Ca- Ala-307),

Alameda County, California. Contrib. Univ. Calif. Archeol. Res. Fac. No. 29,
Appendix G:123-129.

. 1975b. Fish remains from the Stege mounds, Richmond, Contra Costa

County, California. Contrib. Univ. Calif. Archeol. Res. Fac. No. 29, Appendix
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. 1976. Fish remains from an archaeological site at Rancho Carrillo on the Silver

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Frey, Helen. 1974. Preliminary report of a micro-analysis of a San Pedro, California,
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Accepted for publication March 2, 1977.

501 ■ 73
C3 LS6&

NUMBER 290
MARCH 21, 1978

LATE MIOCENE MARINE BIRDS
FROM ORANGE COUNTY, CALIFORNIA

By Hildegarde Howard

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCENC6

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LATE MIOCENE MARINE BIRDS
FROM ORANGE COUNTY, CALIFORNIA

By Hildegarde Howard* 2

Abstract: Bone fragments from five sites in the Late Miocene Monterey
Formation at Laguna Niguel, Orange County, California, represent at least 14
species of marine birds, Gavia brodkorbi n. sp., Diomedea ? calif ornica,
Diomedea sp., Puffinus barnesi n. sp., Oceanodroma sp., Osteodontornis orri
Howard, Morns lompocanus (Miller), Moms magnus n. sp., ?Miosula media
Miller, ?Uria sp., ?Cepphus sp., ?Aethia sp., Fraterculini gen. and sp. indet.,
and Praemancalla wetmorei Howard. The avifauna suggests a slightly younger
phase of the Late Miocene than another avifauna previously reported from the
Monterey Formation in Laguna Hills, three miles northward.

INTRODUCTION

In 1969, during the excavation for the North American Rockwell Building
(now United States General Services Administration Building) on El Lazo
Road in Laguna Niguel, Orange County, California, fossiliferous sands and
siltstones of the Late Miocene Monterey Formation (Clarendonian correlative)
were exposed. Marine mammals and birds were collected in the actual building
excavation and, from 1969 to 1976, in adjacent hillsides within a half-mile
radius of the El Lazo site.

The localities, all of which bear Natural History Museum of Los Angeles
County (LACM) locality numbers, are listed below (numbers in parentheses
indicate the number of avian bones found). Detailed locality descriptions with
reference to the San Juan Capistrano Quadrangle, U.S.G.S. 7.5 minute, 1948
edition, are on file in the Section of Vertebrate Paleontology, LACM.

LACM Loc. 3185 — Aliso Creek. From coarse yellow sand. (3)

LACM Loc. 6901 — El Lazo Road. In laminated gray to white siltstone. (1)
LACM Loc. 6902 — - El Lazo Road. From coarse yellow sands overlying
siltstones described in LACM 6901. (21)

Review Committee for this Contribution
Lawrence G. Barnes
Storrs L. Olson
Stuart Warter

2Chief Curator Emeritus, Natural History Museum of Los Angeles County, 900 Exposi-
tion Boulevard, Los Angeles, California 90007.

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Contributions in Science

No. 290

LACM Loc. 6906 — Site of excavation for North American Rockwell
Building on El Lazo Road. In yellow sands and laminated gray
siltstone. (21)

LACM Loc. 7136 — Moulton Parkway. In phosphatic pebble bed, in a gray
siltstone. (5)

MATERIAL

Fifty-one avian bone fragments were recovered from the Laguna Niguel
localities. These are in the collections of the Natural History Museum of Los
Angeles County (LACM). Thirty-nine are identified and assigned to seven
familes.

Comparative fossil material used in connection with this study is largely in
the LACM collections and includes, in addition to LACM types and referred
specimens, casts of types of Gavia concinna Wetmore 1940; Diomedea califor-
nica Miller 1962; Puffinus conradi Marsh 1870; P. diatomicus Miller 1925; P.
incept or Wetmore 1930; P. mite he Hi Miller 1961; P. priscus Miller 1961; Os-
teodontornis orri Howard 1957; Sula willetti Miller 1925; Morns lompocanus
(Miller 1925); Miosula media Miller 1925; Palaeosula stocktoni (Miller 1935);
and Uria antiqua (Marsh 1870).

In addition, the following material was made available on loan: from the
Museum of Comparative Zoology, Harvard University (MCZ), referred
tibiotarsus (Wetmore 1943) of Diomedea anglica Lydekker 1891; from the
Museum of Paleontology, University of California, Berkeley (UCMP), type
and reverse of type of Miosula media Miller 1925, and figured specimens of
Morus lompocanus (Miller 1925 :pls. 7 and 9); from the United States National
Museum of Natural History (USNM), previously unreported referred
humerus and ulna of Miocepphus mcclungi Wetmore 1940, identified by Storrs
Olson.

Recent skeletal material used for comparison is largely from the LACM
collections, but also includes skeletons of Alcidae obtained on loan from the
Museum of Vertebrate Zoology, University of California, Berkeley (MVZ)
and California State University, Long Beach (CSLB), and a skull of Morus
bassanus lent by Pierce Brodkorb, University of Florida, Gainesville (PB).

HISTORICAL REVIEW

Miller (1925) was the first to document Miocene birds from marine de-
posits in California, naming three species of sulids, a shearwater, a godwit and
an auklet from the Late Miocene diatomaceous shales of the Sisquoc Forma-
tion near Lompoc, Santa Barbara County. Within the next ten years three sites
in Los Angeles County yielded Late Miocene avian fossils: the Modelo For-
mation at Calabasas (Miller 1929), and the Monterey Formation at Lomita and
San Pedro (Miller 1935). Also, in the same decade (1925-1935), the first avian
bones from the Middle Miocene (Round Mountain Silt) Sharktooth Hill

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Miocene Marine Birds

3

Bonebed were recorded (Wetmore 1930). By the end of 1935, 11 species had
been described.

No further marine birds from the California Miocene were added until the
1950’s when a flagstone quarry in the Monterey Formation in Tepusquet Can-
yon, Santa Barbara County, yielded two avian skeletons. Both were described
under extinct families (Howard 1957a and 1957b). Later excavations in the San
Fernando Valley and at El Sereno, Los Angeles County, yielded additional
records of some of the previously described species and added a new sulid
(Howard 1958, and Howard and White 1962). Miller (1951) described a storm
petrel during this decade, from the Capistrano Formation near San Juan Capis-
trano, Orange County. The age of the deposit was given as Middle Miocene,
but is now considered to be either Late Miocene or Early Pliocene (Fife
1974:19).

The 1960’s added two Miocene bird localities in California: the Jewett
Sand at Pyramid Hill, Kern County (Early Miocene) yielded a single bone
described in a new family related to the cormorants (Howard 1969), and a large
collection from the Monterey Formation at Leisure World, Laguna Hills,
Orange County added five new species (Howard 1966a and 1968). Also, within
the last 16 years, five additional species have been described from the
Sharktooth Hill Bonebed (Miller 1961 and 1962; Howard 1966b; Warter 1976).

In seven of the 1 1 areas in California from which Miocene marine birds
have been previously obtained, the specimens occur as partial skeletons or
skeletal impressions on slabs of shale. Such specimens include the types of 12
of the 26 recorded species. While these specimens provide information on the
proportions of the birds involved, the finer details of structure are usually not
clearly preserved. Consequently it becomes difficult to compare these speci-
mens with the isolated, mineralized bone fragments obtained from localities
such as Sharktooth Hill, Pyramid Hill, Laguna Hills Leisure World, or the area
discussed herein.

In the following list of species from previously recorded marine Miocene
sites in California, those based on partial skeletons in shale slabs are marked
with an asterisk.

Procellariiformes

Diomedeidae: Diomedea californica Miller 1962; D. milleri Howard 1966.

Procellariidae: *Puffmus diatomic us Miller 1925; P. inceptor Wetmore
1930; P. mitchelli Miller 1961; P. prisons Miller 1961; P. calhouni
Howard 1968; Fulmar us hammeri Howard 1968.

Hydrobatidae: *Oceanodroma hubbsi Miller 1951.

Pelecaniformes

Pseudodontornithidae; *Osteodontornis orri Howard 1957.

Sulidae: *Sula willetti Miller 1925; *Sula polili Howard 1958; *Morus
lompocanus (Miller 1925); M. vagabundus Wetmore 1930;
*Palaeosula Stockton i (Miller 1935); *Miosula media Miller 1925.

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Phalacrocoracidae: *Phcilacrocorax femoralis Miller 1929.

Plotopteridae: Plotopterum joaqainensis Howard 1969.

Anseriformes

Anatidae: Presbychen abavus Wetmore 1930.

Falconiformes

Pandionidae: Pandion homalopteron Warter 1976.

Charadriiformes

Scolopacidae: *Limosa vanrossemi Miller 1925.

Alcidae: Aethia rossmoori Howard 1968; *Cerorhinca dubia Miller 1925;
Alcodes idnulus Howard 1968; Praemancalla Icigunensis Howard
1966.

Passeriformes

Palaeoscinidae: *PaIaeoscinis turdirostris Howard 1957.

The species represented at Laguna Niguel bring the total for the Alcidae to
five identified species, and the totals for the Procellariidae and the Sulidae to
seven species each. The Order Gaviiformes (Gaviidae, 1 species) is added to
the California Miocene list.

SYSTEMATICS
Order GAVIIFORMES

Family Gaviidae — Loons
Genus Gavia Forster 1788
Gavia brodkorbi new species

Figure 1 a, b

Holotype. — Complete left ulna, LACM 31173, collected by Marion J.
Bohreer, 1969, from locality LACM 6906.

Diagnosis. — Ulna relatively short and stout; proximally, attachment for
anterior articular ligament short and broad (roughly triangular), and promi-
nently set off from shaft, with brachial impression deeply rimming its palmar
edge; distal tip of external cotyla bent toward shaft, with short scar running
mediad directly beneath, confining small radial impression; distally, large car-
pal tuberosity jutting abruptly from shaft.

Measurements. — Greatest length 81.0 mm, breadth across proximal
cotylae 9.4 mm, breadth of shaft at middle 4.9 mm, greatest breadth of distal
and (through carpal tuberosity) 1 1.3 mm, depth of distal end through external
crest of trochlea 7.5 mm, length of attachment for anterior articular ligament
4.4 mm, breadth of same 4.0 mm.

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Miocene Marine Birds

5

Etymology. — The species is named in honor of Pierce Brodkorb in recog-
nition of his many contributions to Paleornithology, including a review of fossil
loons.

Discussion. — The fossil ulna is 25 mm (23.6%) shorter than the minimum
for this element in four LACM specimens of the Red-throated Loon, Gavia
stellata (Pontoppidan 1763), but is relatively stouter. The proximal radial im-
pression is more confined than in G. stellata, G. pacifica (Lawrence 1848), or
G. immer (Brunnich 1764). Distally, the carpal tuberosity is more square in
outline than in these Recent loons. The short, broad attachment of the anterior
articular ligament, also, is distinct (the attachment is longer and more oval in
the Recent species).

A photograph of an ulna of Colymboides minutus Milne-Edwards 1867
from the Early Miocene of France, illustrated by Storer (1956, Fig. 1, g) shows
the attachment of the anterior articular ligament to be broad and short. Storer,
however, notes that, unlike the ulna of all Recent loons, this element of Co-
lymboides lacks the groove bordering the attachment posteriorly. This groove
is present in G. brodkorbi. Furthermore, the carpal tuberosity in Colymboides
is less abruptly projected than in G. brodkorbi or in any of the Recent loons.

The only previous Miocene record of the genus Gavia was based on a
poorly preserved tibiotarsus from the Calvert Formation, Maryland, cited by
Wetmore (1941) as Gavia sp. Four species have been described from the
Pliocene: Gavia portisi (Regalia 1902), Middle Pliocene of Italy; Gavia con-
cinna Wetmore 1940, Early Pliocene of Florida and Middle and Later Pliocene
of California; Gavia palaeodytes Wetmore 1943, Early Pliocene of Florida;
Gavia howardae Brodkorb 1953, Late Pliocene of California.

According to Brodkorb (1953), who reviewed these species, Gavia portisi
is known only from a cervical vertebra that is nearly as large as that of G.
immer. The type of G. concinna is an ulna much larger than that of G. brod-
korbi (breadth across proximal cotylae 11.6 mm), and is further distinguished
by a longer attachment for the anterior articular ligament. G. palaeodytes is
known from coracoid, humerus and femur, all equal to, or slightly larger than
comparable specimens of G. stellata, hence larger than would be expected for
G. brodkorbi.

Gavia howardae was described from an incomplete humerus with two
additional humeral specimens referred (all LACM). The smallest referred
humerus provides a measurement of length (from distal end to distal tip of
deltoid crest) of 91.5 mm, which is 14% less than the minimum for this same
measurement in G. stellata (106.5 mm). The type of G. howardae is incom-
plete, but appears to have been longer than the referred specimen (possibly
within 7 mm of the minimum for G. stellata). Relative breadth is difficult to
determine in these incomplete specimens. They appear, however, to be of less
stocky proportions than the ulna of G. brodkorbi. Qualitatively there is little on
which to base comparison of the humerus of G. howardae with the ulna of G.
brodkorbi. However, the long, narrow attachment for the anterior articular

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ligament on the humerus of G. howardae closely resembles the condition found
in G. stellata and is unlikely to correspond with the unusually short, broad
attachment for this ligament on the ulna of G. brodkorbi.

Order PROCELLARIIFORMES
Family Diomedeidae — Albatrosses
Genus Diomedea Linnaeus 1758
Diomedea ? calif ornica Miller 1962

Figure 2 c

Referred material. — Distal end of tibiotarsus, LACM 37629, from lo-
cality LACM 6906.

Discussion. — Four species of fossil albatrosses have been previously
named: D. californica Miller 1962 and D. milleri Howard 1966b, from the
Middle Miocene of Sharktooth Hill, California; D. thyridata Wilkinson 1969,
from the Late Miocene of Australia; and D. anglica Lydekker 1891, from the
Pliocene of England (type) and Florida (specimen referred by Wetmore 1943).

D. milleri was described from an ulna with referred tarsometatarsus, both
of which are smaller than comparable elements ofD. nigripes Audubon 1839.
D. thyridata, described from a rostrum, is likened in characters and size (Wil-
kinson 1969) to D. melanophris Temminck 1828. It would appear, therefore,
that both D. milleri and D. thyridata were species whose size range was below
that possible for the species represented by the tibiotarsus in the present collec-
tion.

D. californica and D. anglica, both described from the tarsometatarsus,
were larger species than either D. milleri or D. thyridata. The referred speci-
men of D. anglica is a distal end of tibiotarsus. This specimen (MCZ 2328) was
made available for the present study (Fig. 2, B). In distal breadth LACM 37629
from Laguna Niguel is only slightly larger than MCZ 2328. It differs from the
latter, however, in less depression of the supratendinal bridge, and in having a
well-developed, papilla-like internal ligamental attachment. In both of these
characters LACM 37269 resembles D. exulans Linnaeus 1758, whereas MCZ
2328 more closely resembles D. albatrus Pallas 1769, in which the bridge is
more depressed and the ligamental attachment is only a scar. Both fossil
specimens differ from D. exulans in more horizontal position of the lower
opening of the tendinal canal, but in MCZ 2328 the opening is more restricted
in lateral extent than in LACM 37629.

Comparison of the Laguna Niguel specimen with D. californica rests en-
tirely on size, as no tibiotarsus assignable to this species has yet been forthcom-
ing from the Sharktooth Hill Bonebed. A second tarsometatarsus (LACM
18203) from that locality is, however, now at hand. This is slightly larger, but
otherwise similar to the holotype. The distal breadths in the two tarsometatarsi
ofZ). californica are 92.3% (holotype) and 96.4% (LACM 18203, referred) of

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this dimension in a tarsometatarsus (LACM Bi230) of D. exulans. Compared
with the tibiotarsus of the same specimen of D. exulans, the distal breadth of
tibiotarsus LACM 37629 from Laguna Niguel is 96.7% (Table I). It appears
reasonable, therefore, to assign this specimen to D. californica. However,
being unable to compare it with a tibiotarsus from the type locality of D.
californica, the assignment is tentative.

Diomedea sp. indeterminate
Figure 2 a, d

Referred material. — Proximal section of humerus, LACM 58544, from
locality LACM 6902, and distal end of radius, LACM 31172, from locality
LACM 6906.

Discussion. — These poorly preserved wing bones are notably smaller
relative to those of D. exulans than is the case with the tarsometatarsi of D.
californica or tibiotarsus LACM 37629. They are somewhat smaller, also, than
a distal end of humerus from the Sharktooth Hill Bonebed previously referred
to D. californica (Howard 1966b) (see Table I).

The incomplete humerus lacks the tip of the internal tuberosity, the bicipi-
tal crest and a large portion of the deltoid crest. It resembles the humerus ofD.
exulans in the broad curvature of the anconal face of the shaft, but the area
immediately below the head is less depressed. On the palmar surface, the
enlarged distal tip of the deltoid crest resembles the condition in D. exulans,
but the crest is much shorter in the fossil. The radius is too poorly preserved
for analysis, but provides a measurement of distal breadth (Table I).

Family Procellariidae — Shearwaters

Genus Puffinus Brisson 1790

Four wing bones are referable to the genus Puffinus. Thirteen species of
middle to late Tertiary shearwaters of this genus have been described from
Europe and North America, eight of which are from the west coast (see Brod-
korb 1963b, and Howard 1968 and 1971).

After consideration of these species, and careful comparison with speci-
mens of all those from California, I find it necessary to add still another
species, which I refer to the subgenus Puffinus.

Puffinus barnesi new species
Figure 1 e, f

Holotype. — Left humerus lacking proximal end, LACM 42652, collected
by W. Earl Calhoun, July 1969, from locality LACM 6906.

Diagnosis. — Humerus with shaft laterally compressed above distal end.

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TABLE 1

Comparison of Fossil and Recent Specimens of Diomedea

Ratio Fossils to Recent

Measurements in Millimeters (In Per Cent)

D. calif -
ornica*

D. sp.*

*D. ang-D. exu-
lica*** Ians****

D. calif-
ornica*

D. sp.**

D. ang-
lica***

Tarsometatarsus

Distal breadth

20.6-21.5

19.3 22.3

92.4- 96.4

86.5

Breadth shaft

9.0-10.0

9.1

96.8-109.0

Proximal breadth

20.7 23.2

89.1

Tibiotarsus

Distal breadth

20.6

20.2 21.3

96.7

94.8

Breadth shaft

10.0

9.5 10.0

100.0

95.0

Humerus

Proximal breadth

39.2

48.5

80.8

Distance head to

end deltoid crest

64.5

.... 83.7

77.0

Distal breadth

27.5

31.6

87.0

Radius

Distal breadth

11.7

14.6

80.0

*Type and referred specimens from Sharktooth Hill
**Specimens from Laguna Niguel (tibiotarsus referred D. ?californica)
***Type tarsometatarsus, referred tibiotarsus
****LACM no. Bi230

but slightly rounded in contour; internal side of distal end relatively short in
anconopalmar dimension, and anconal tip swollen laterally; impression of
brachialis anticus small, and short in proximo-distal dimension; ectepicondylar
process situated relatively near to distal end; attachment of anterior articular
ligament turned slightly laterally rather than facing directly palmad.

Measurements. — Length from distal end to distal tip of deltoid crest 67.0
mm (estimated total length 80 mm), breadth of distal condyles 7.7 mm, depth of
internal side of distal end 8.4 mm, distance from distal surface of condyle to
proximal edge of ectepicondylar process 9.5 mm, shaft dimensions near distal
end 3.5 mm in breadth, 5.7 mm in depth, shaft dimensions (middle) 3.9 mm in
breadth, 5.9 mm in depth.

Etymology. — The species is named for Lawrence G. Barnes in recogni-
tion of his paleontological studies of the marine vertebrates of the west coast.

Discussion. — The holotype of P. barnesi is comparable in general size to
the humerus of Puffinus opisthomelas Coues 1864. Although the shaft is com-
pressed laterally as in that Recent species, it is slightly more rounded and less
bladelike in the fossil.

With the exception of Puffinus tedfordi Howard 1971, from the Almejas

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9

Formation of Cedros Island, Baja California, Mexico, and P. arvernensis
Milne-Edwards 1871, from the Early Miocene of France, all previously de-
scribed Tertiary representatives of this genus are known from the humerus.
Shufeldt (1896) referred a humerus to P. arvernensis, but according to Storrs
Olson (personal communication), who has examined the specimen, it comes
from a Pleistocene locality and should be assigned to the Recent species Puf-
finus puffinus (Brunnich 1764). The holotypes of both/5, tedfordi and P. arver-
nensis are tarsometatarsi. In both species, this element suggests a stouter bird
than is represented by the wing of P. barnesi.

Judging from the descriptions given by Milne-Edwards (1874), the type
humeri of his species Puffinus aquitanicus and P. antiquus, from the Middle
Miocene of France, both exceed P. barnesi in size. Also, as indicated by a cast
of the type of P. conradi Marsh 1870 (LACM C688), that species was markedly
larger than P. barnesi. At the opposite extreme, Brodkorb’s (1963a: 161) meas-
urements of the type humerus of his species/5, micraulax, from the Hawthorne
Formation, Early Miocene of Florida, show that species to be notably smaller
than P. barnesi.

Original material or casts of all the California species of Tertiary Puffinus
have been examined, including topotypical specimens of humeri referable to P.
mitchelli Miller 1961 (LACM 17500) and P. priscus Miller 1961 (LACM 17502
and LACM 18140) recovered since the last report on the avifauna of the Mid-
dle Miocene of Sharktooth Hill (Howard 1966b).

Of the California species, the humeri of P.felthami Howard 1949 (Middle
Pliocene of Orange County) and P. mitchelli are larger than P. barnesi. P.
inceptor Wetmore 1930 (Middle Miocene, Sharktooth Hill) agrees in some
dimensions, but the marked medial thrust of the internal condyle and the
greater anconopalmar dimension of the internal side of the distal end are dis-
tinctive characters of P. inceptor. Also, the brachial impression in that species
is “more distally developed. In P. calhouni Howard 1968 (Late Miocene,
Orange County) and P. priscus the shaft is more compressed and bladelike
than in P. barnesi. The ratio of breadth to depth of shaft near the distal end in
P. barnesi is 61%, in P. calhouni 50%, and in P. priscus 48-53%.

An excellent relief cast (LACM C692) from the holotype skeletal impres-
sion of P. diatomicus Miller 1925 (Late Miocene, Lompoc) clearly reveals
characters of the palmar aspect of the distal end of the humerus. The impres-
sion of the brachialis anticus is more distally extended than in P. barnesi and
the ectepicondylar process is placed higher above the distal end (11.6 mm from
distal surface of condyle to proximal tip of process).

Most closely approaching P. barnesi in general size are the paratype and
referred humeri of P. kanakoffi Howard 1949 (LACM specimens), from the
San Diego Formation. This Pliocene species is distinguished, however, by a
longer brachial impression, less rounded shaft (though less compressed than in
P. priscus), less inflated anconal tip of the internal condyle, and more palmad-
facing attachment for the anterior articular ligament.

Referred material. — A left ulna, lacking the olecranon (LACM 42654)

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Figure 1. A, B, Gavia brodkorbi n. sp. holotype ulna, LACM 31 173, internal and palmar
views. C, D, Morus lompocanus Miller referred carpometacarpus, LACM 77697 , inter-
nal and external views. E, F, Puffinus barnesi n. sp. holotype humerus, LACM 42652,
palmar and internal views. X 1

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Figure 2. A and D, Diomedea sp. indet., proximal portion of humerus, LACM 58544,
anconal and palmar views. B, Diomedea anglica Lydekker, referred distal end of
tibiotarsus from Pierce, Florida, MCZ 2328, anterior view. C, Diomedea ?californica
Miller, referred distal end of tibiotarsus, LACM 37629, anterior view. X 1

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Figure 3. A and D, Osteodontornis orri Howard, referred proximal half of left car-
pometacarpus, LACM 53906, posterior and internal views. B, C, Morns magnus n. sp.
holotype carpometacarpus, LACM 37632, internal and external views. X 1

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Figure 4. A, B, Morus magnus n. sp., referred rostrum, LACM 77696, ventral and
dorsal views, C, D, Sulidae sp. indet., rostrum, LACM 73565, dorsal and ventral views.
X 1

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from the type locality, LACM 6906. Although smaller than would be expected
for the wing of the same individual as the holotype, this specimen (like the
humerus) falls within the size range ofP. opisthomelas . (ulnar length, 68.3-76.3
mm), though near the minimum; the holotype humerus is near the maximum
size ofP. opisthomelas. The few characters discernible in the poorly preserved
ulna are: carpal process as in P. opisthomelas, shorter than in a topotype ulna
of P. kanakoffi (LACM 2821, previously unreported), and more abruptly pro-
jected; external cotyla not prominently projected palmad; ridge present from
external cotyla to shaft; attachment for anterior articular ligament seemingly
more protruding then in P. opisthomelas; but the area is broken.

Measurements of ulna: length to intercotylar ridge 70.8 mm, breadth of
proximal end 6.5 mm, greatest breadth of distal end 6.4 mm, depth through
external crest of trochlea 5.3 mm, shaft dimensions (middle) 3.5 mm x 5.3 mm.

Another poorly preserved fragment of a distal end of an ulna (LACM
52748) from locality LACM 7136, has a distal breadth of approximately 6.3
mm. A wing phalanx (digit 2, phalanx 1), LACM 53925 from locality LACM
3185, measures 20.4 mm in length, which is within the size range of this element
of P. opisthomelas. These two specimens are tentatively assigned to P. bar-
nesi.

Family Hydrobatidae — Storm Petrels
Genus Oceanodroma Reinchenbach 1852
Oceanodroma sp. indeterminate

Referred material. — A single incomplete right tarsometatarus, LACM
42659, from locality LACM 6902, represents this family and genus.

Discussion. — The specimen resembles this element in O. melania
(Bonaparte 1854), but is slightly longer. The length from distal end to proximal
tip of hypotarsus is 31.5 mm. The distal end appears to be narrower than in O.
melania, but the preservation is such that accurate measurement is impossible.

The only previously recorded Tertiary storm petrel is O. hubbsi Miller
1951, from the Capistrano Formation (Late Miocene-Early Pliocene), Orange
County, a few miles south of the Laguna Niguel localities. The tarsometatarsus
is represented in the type, partial skeleton, of this species preserved in shale.
Miller (1951) gives the length of this element as 22.4 mm, markedly less than
even the incomplete measurement possible on the specimen now at hand.

Order PELECANIFORMES
Family Pseudodontornithidae — Bony-toothed Birds
Genus Osteodontornis Howard 1957
Osteodontornis orri Howard 1957
Figure 3 a, d

Referred material. — Lower jaw fragment, LACM 22444, and car-
pometarcarpus, LACM 53906, with portion of proximal end and shaft, both

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15

from locality LACM 6902; portion of lower jaw, LACM 42656, with a single
“tooth,” from locality LACM 7136.

Discussion. — The jaw fragments resemble previously recorded speci-
mens of Osteodontornis (Howard 1957a, and Howard and White 1962), al-
though they seem somewhat smaller than the type of O. orri.

The fragment of carpometacarpus, however, conforms in size to the left
carpometacarpus on the type skeleton, a cast of which is at hand. In fact, the
specimen from locality 6902 fits exactly into the impression of the left car-
pometarcarpus on type Slab No. 1 (LACM C703, Block No. 1), in which the
large pisiform process has left a deep depression. The present specimen not
only clarifies that this depression, was, indeed, made by the pisiform process,
but also reveals other characters only suggested in the type, namely, the length
of the process of metacarpal 1 and the great compression of metacarpal 3(M3)
to metacarpal 2(M2). The very thin M3 is pressed against M2 even distal to the
proximal metacarpal symphysis, and the symphysis itself extends almost to the
level of the distal tip of the process of ML

Metacarpal 1 is broken in both the type and LACM 53906, but the area of
its attachment to the element as a whole is indicated in the type and clarified in
the broken edges of the process on the carpometacarpus from Laguna Niguel.
The length of Ml measured on LACM 53906, is 62.2 mm, approximately
one-fourth the total length of the carpometacarpus as seen in the type (252
mm). Enough of the proximal end of LACM 53906 is preserved to obtain an
approximate breadth of the proximal trochlea (15.2 mm). The antero-posterior
dimension of the trochlea cannot be measured, but it is obvious that the post-
erior portion is short in distal extent. Shaft breadths of M2 and M3 are 1 1 .0 mm
and 4.0 mm respectively. The depth through the compressed M2 and M3 is
16.4 mm. The incomplete specimen measures 126.4 mm from trochlea to bro-
ken end of M2.

Except for a prominent pneumatic foramen above the pisiform process,
which occurs in most sulids, there is nothing about this highly compressed
carpometacarpus to relate it to the Pelecaniformes. I have previously con-
tended (Howard 1957a) that the bony-toothed birds represent a distinct order,
Odontopterigiformes.

Family Sulidae — Boobies and Gannets

The family Sulidae is the best represented family in the Laguna Niguel
collection and the one which has presented the greatest difficulty in identifica-
tion. Fifteen middle and late Tertiary sulids have been previously described
from North America and Europe (see Brodkorb 1 963b:257-26 1 ), eight of which
are from California (six Miocene, two Pliocene). Five of the species are based
on partial skeletons in shale slabs, the others on individual bones involving four
different incomplete skeletal elements. A complete review of the known fossils
of this family is greatly to be desired. This is a task that some energetic young
paleontologist may profitably undertake.

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Sixteen bones in the present collection are assignable to at least three
species. In spite of the difficulty experienced in correlating the previously
described species, I feel justified in describing one new species and in assigning
several specimens to one previously described.

Genus Morus Viellot 1816
Morns lompocanus (Miller 1925)

Figure 1 c, d

Referred material. — Carpometarcarpus, LACM 37634; distal end of
ulna, LACM 37636; and distal end of femur, LACM 37633; all from locality
LACM 6902; carpometacarpus, LACM 77697, from locality 6901; proximal
end of tarsometatarsus, LACM 32428 from locality LACM 6906; proximal end
of tarsometatarsus, LACM 42657, from locality LACM 7136; distal end of
tibiotarsus, LACM 52217, from locality LACM 3185.

Discussion. — Morus lompocanus was described (Miller 1925) from the
impression of a partial skeleton in a slab of diatomaceous shale from Lompoc,
California. Although the holotype (UCMP 26544) was the only specimen de-
scribed in the text, two other partial skeletons from the same site were so
named and illustrated (op. cit.:pl. 7B and pi. 9). These referred specimens
(UCMP 117309 and UCMP 115855), both bearing Miller’s identification, as
well as a cast (in relief) of the holotype (LACM C697) are at hand. On the basis
of comparison of size with these specimens, the fossils from Laguna Niguel are
referred to M. lompocanus.

The two carpometacarpi measure 94.7 mm (LACM 37634) and 95.7 mm
(LACM 77697) in length. By comparison, a carpometacarpus (LACM Bi 1764)
of a female M. bassanus (Linnaeus 1758) is 90.0 mm, and Miller (1935:78)
records another of this Recent species at 94 mm. The referred carpometacar-
pus of M. lompocanus (UCMP 115855) is 96.6 mm. Other fossil sulids in which
this dimension is known are: Palaeosula stocktoni (Miller 1935), 102 mm;
Miosula media Miller 1925, 81 mm; Sula willetti Miller 1925, 70 mm; S. pohli
Howard 1958, 69 mm; Microsula avita (Wetmore 1938), 75 mm.

Carpometacarpus LACM 77697, the better preserved of the two Laguna
Niguel carpometacarpi (Fig. 1 C, D), is similar in the shape of the process of
Ml to Morus bassanus. But where the small foramina occur in the modern
species, there is a deep, slit-like foramen — one anterior to the pisiform pro-
cess, and one in a similar position at the base of Ml on the external side of the
proximal end. The fossil also resembles Morus in the absence of pneumatic
foramina at the posterior edge of the trochlea. Details of qualitative characters
are not discernible on the Lompoc specimens.

Ulna LACM 37636 resemble Morus rather than Sula in the pneumaticity
of the palmar face of the carpal process, and the size of the process. It is of the
same distal breadth (10.5 mm) as the ulna of M. bassanus LACM Bi 1764.
Measurements of ulnar breadth cannot be made on M. lompocanus, but the
Laguna Niguel specimen corresponds favorably with the impression of the
element on Lompoc specimen UCMP 115855.

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17

Femur LACM 37633 conforms in proportions to a raised mold of the
element made from the type impression of M. lompocanus. Similar dimensions
measured on the two specimens are identical: breadth across anterior face of
distal end 13.9 mm, breadth of shaft 7.0 mm. M. bassanus LACM Bi 1764 is
larger in these dimensions (14.2 mm and 7.5 mm, respectively).

Tibiotarsus LACM 52217 conforms in size withM. lompocanus specimen
UCMP 1 15855 from Lompoc. The bridge is less vertical in position than in M.
bassanus, the upper edge being tipped posteriorly. This is true, as well, of the
type tibiotarsus of Miosula recentior Howard 1949 (LACM 2117) from the
Pliocene of San Diego, California. But the latter is distinguished from the
specimen now at hand by less vertically oriented condyles. Measurements of
LACM 52217 are: breadth of distal end 13.7 mm, depth of distal end 12.4 mm,
ratio of depth to breath 90.5%. The same dimensions in M. bassanus LACM Bi
1764 are, breadth 13.0 mm, depth 12.0 mm, ratio depth to breadth 92%.

The two proximal ends of tarsometatarsi (LACM 32428 and LACM
42657) measure 12.8 mm and 12.9 mm in proximal breadth, respectively. A
mold of the posterior surface of this element on the type specimen of M.
lompocanus measures 12.8 mm proximally. The impression visible on Lompoc
specimen UCMP 115855 measures 13.5 mm. M. bassanus and a mold of the
tarsometatarsus in the type of Miosula media Miller 1925 are broader (14.5 mm
and 14.1 mm, respectively).

Morus magnus new species
Figure 3 b, c; 4 a, b

Holotype. — Nearly complete left carpometacarpus, LACM 37632, col-
lected by Marion J. Bohreer, 1969, from LACM locality 6906.

Diagnosis. — More than 20% longer than female specimens of this ele-
ment of Morus bassanus. Area above pisiform process depressed, with small
pneumatic orifice. Externally, a deep, slit-like depression at base of process of
metacarpal 1.

Measurements. — Greatest length 116.1 mm, breadth proximal trochlea
11 mm (approximately), breadth of shaft 9.6 mm.

Etymology . — The species name magnus (Latin, great) refers to the large
size of the skeletal elements described.

Discussion. — Owing to the poor preservation of the holotype car-
pometacarpus, size is the outstanding distinguishing character. Of previously
described sulids, Palaeosula stocktoni most nearly approaches M. magnus in
length of this element (102 mm), but is still 13% shorter. Characters of the
carpometacarpus other than length are not clearly discernible on the type slab
of P. stocktoni (cast, LACM C743). It has been shown, however (Howard
1958), that at least the humerus of Palaeosula is markedly distinct qualitatively
from either Morus or Sula. M. magnus , on the other hand, resembles the living
sulids.

From the elements known, none of the other middle to late Tertiary fossil
sulids gives evidence of approaching M. magnus in size.

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No. 290

Referred material. — Distal end of humerus, LACM 32430, and shaft of
femur, LACM 37628, both from the type locality, LACM 6906, collected by
Bohreer in 1969; and rostrum, LACM 77696, from locality LACM 6902, col-
lected by Jennifer Whistler, June, 1975.

The humerus resembles Morus in contrast to Sula in the absence of the
deep pneumatic foramen undercutting the external side of the olecranal fossa.
The flat surface of the attachment for the anterior articular ligament, and its
length relative to the distal breadth of the element is also similar to Morus. The
attachment, however, projects slightly more palmad at its proximal end than in
M. bassanus, and, in this respect, resembles Sula. In breadth the element is
22% greater than in M. bassanus.

The fragmentary femur is 18% broader than in M. bassanus in least
breadth of shaft, and appears to expand to even greater relative breadth farther
distad. It is difficult to be certain of accurately reproducing the same measure-
ment in the two specimens as the distal condyles are lacking in the fossil.

The rostrum resembles Sula in the broad arch of the nasals, but bears
resemblance to Morus in the depression of the dorsal contour anterior to the
hinge. This depressed area, however, is shorter than in M. bassanus. In
greatest breadth, it exceeds the rostrum of M. bassanus by 31% (see Table 2).

?Miosula media Miller 1925

Referred material. — Tarsometatarsus lacking the proximal end, LACM
32431, from locality LACM 6906.

Discussion. — This specimen is heavier of shaft than the two proximal
fragments of tarsometatarsus referred to Morus lompocanus. In breadth of
distal end (17.6 mm) it is slightly larger than an incomplete mold of the tar-
sometatarsus made from the type skeletal impression of Miosula media (17.3
mm, approx.). Although neither the mold nor LACM 32431 is complete, both
suggest a tarsometatsus that is straighter of shaft than in Morus, but having the
slightly raised internal trochlea characteristic of that genus as distinguished
from Sula.

SULIDAE, Spp. INDETERMINATE

Referred material. — A rostrum, LACM 73565, a proximal end of radius,
LACM 58551, and a distal end of tibiotarsus, LACM 57834, all from locality
LACM 6902; and a rostrum, LACM 37614, from locality LACM 6906.

Discussion. — Rostrum LACM 73565 (Fig. 4 C, D) bears strong re-
semblance to the cormorants in dorsal aspect, having deep lateral grooves
setting off a narrow nasal process. There are, however, no dorsal foramina
such as occur in Phalacrocorax. In palatal view, the resemblance is closer to
the sulids. The palatines in the maxillo-palatine area are swollen, not flat, and
are bordered laterally with wide, deep grooves. Posteriorly the maxillo-
palatine area slants smoothly upward and bears very little perforation. In the
slope of the area the specimen resembles Morus, in perforation it resembles

1978

Miocene Marine Birds

19

TABLE 2

Measurements (in millimeters) of Morus magnus and M. bassanus

M. magnus M. bassanus

(LACM Bi 1764)

Carpometacarpus

Greatest length

116.1

90.0

Breadth proximal trochlea

1 1 .0 approx.

9.9

Breadth middle of shaft

9.6

7.4

Height process Ml

16.0

12.0

Humerus

Greatest breadth distal end

29.2

24.1

Depth distal end (externally)

17.0

14.4

Breadth of shaft immediately
proximal to attachment for

anterior articular ligament

22.8

17.0

Femur

Least breadth shaft

8.9

7.1

Depth of shaft

9.6

7.5

Rostrum

Breadth of frontonasal hinge

34.8

25.4-26.5*

Greatest depth

25.3

15.4-16.7*

Breadth nasal process

31.4

18.3-19.5*

Length

138.0 (estimate)

101 .7-106.5

*Maximum measurements from rostrum PB 16291, minimum from LACM Bi 1765.

both Phalacrocorax and Moms, but not Sulci. This area in Sula rises more
abruptly and is well perforated. The lateral pneumatic openings are much
reduced in comparison to those found in the cormorants, and compare more
favorably with those of Moms. Measurements of LACM 73565: greatest
breadth 22.2 mm (approximately), greatest depth, 15:1 mm, breadth nasal pro-
cess 5.2 mm.

In view of the fact that Miller (1925) noted certain characteristics of
Miosula media that are cormorant-like, it is possible that this rostrum may
represent that species.

The second rostrum (LACM 37614) is typically sulid in all aspects. It is
more massive than LACM 73565, but smaller than LACM 77696 referred to
Moms magnus. Measurements: greatest breadth 24.0 mm, greatest depth 19.2
mm, breadth nasal process 21.8 mm.

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Contributions in Science

No. 290

The radius resembles Morns bass anus in general conformation and size,
but is poorly preserved. The tibiotarsus is similar in characters of the distal end
to LACM 52217 from Laguna Niguel, assigned to Moms lompocanus. The
specimen is eroded so that measurements cannot be made precisely. It ap-
pears, however, to be narrower but relatively deeper than in LACM 52217,
breadth of distal end 12.0 mm (approx.), depth of distal end 1 1 .5 mm (approx.);
ratio of depth to breadth 96%.

Order CHARADRIIFORMES
Family Alcidae — Auklike Birds

Discussion. — In addition to the six bones of Praemancalla wetmorei
(subfamily Mancallinae) described earlier (Howard 1976) from localities
LACM 6902, 6906 and 3185, the alcids from Laguna Niguel are represented by
five fragments assignable to the Alcinae. In view of the fact that comprehen-
sive studies of Tertiary alcids are under way by Storrs Olson at the United
States National Museum of Natural History (Atlantic avifauna) and G. Victor
Morejohn at California State University, San Jose (Pacific avifauna), I have
refrained from attempting to name these poorly preserved specimens.

Five middle to late Tertiary alcines have been previously described from
the west coast: Aethia rossmoori Howard 1968, and Cerorhinca dubia Miller
1925, from the Late Miocene; Brachyramphus pliocenus Howard 1949,
Ptychoramphus tenuis Miller and Bowman 1958, and Cerorhinca minor How-
ard 1971, from the Middle to Late Pliocene. East coast species are Uria anti-
qua (Marsh 1870), and Miocepphus mcclungi Wetmore 1940, from the Middle
Miocene; and Australca grandis Brodkorb 1955, from the Middle Pliocene. A
single species, Uria ausonia Portis 1887, is recorded from the Middle Pliocene
of Italy (see Brodkorb 1967; and Howard 1968 and 1971). There is no indication
that any of these species is represented by the material from the Laguna Niguel
sites.

Genus Uria Brisson 1760
?Uria sp.

Referred material. — An incomplete proximal end of humerus, LACM
52018, from locality LACM 6902.

Discussion. — This poorly preserved specimen resembles the humerus in
the murres in the long, oval pectoral scar, and prominent head widening to-
wards the internal side and overhanging a broadly depressed tricipital area
between the pectoral scar and the internal tuberosity. Both the deltoid and the
bicipital crests are incomplete. That which remains of the bicipital surface is
prominently raised proximally and bordered medially below by a deep groove;
the bicipital furrow is a deep notch. These characters are most closely matched
by humeri of Recent U. lomvia (Linnaeus 1758).

Comparisons were made with a cast of the type of Uria antiqua, provided
through the courtesy of Storrs Olson, United States National Museum of

1978

Miocene Marine Birds

21

Natural History. Olson (personal correspondence) now refers this species to
the genus Australca Brodkorb 1965. The cast is not only larger, but is much
flatter in the bicipital area than in the Laguna Niguel specimen.

Genus Cepphus Pallas 1769
?Cepphus sp.

Referred material. — A proximal end of ulna with incomplete olecranon,
LACM 47045, from locality LACM 6906.

Discussion. — This specimen resembles the ulna of the Pigeon Guillemot,
Cepphus columba Pallas 1811, in the rounded shaft lacking a distinct keel
anconally, and in having the brachial impression bordering the attachment for
the anterior articular ligament and extending farther proximally than in most
other genera of alcines. However, the brachial impression is broader in the
fossil, and more deeply undercuts the attachment for the anterior articular
ligament than in the Recent species. Also the attachment for the ligament is
more prominent and more square in outline.

The possibility that this specimen might represent the genus Miocepphus
prompted me to contact Storrs Olson. He reports (personal correspondence)
that “ Miocepphus mcclungi is one of the commoner birds in the Calvert For-
mation of Maryland and Virginia and I have many specimens that have not
been reported on in the literature.” He provided me with the loan of a complete
ulna and humerus which he said “certainly pertain to this species.” In the ulna
(USNM 237219) the attachment for the anterior articular ligament is less prom-
inent and more elongated than in LACM 47045, the brachial impression is
narrower, and the shaft more compressed. LACM 47045 is not related to
Miocepphus, which accords with Olson’s observation that ” Miocepphus is not
related0 to Cepphus ” but to the “ Alca-Uria group of Atlantic alcids.”

Genus Aethia Merrem 1788
?Aethia sp.

Referred material. — Incomplete humerus, LACM 37686, from locality
LACM 6906.

Discussion. — LACM 37686 is the smallest of the alcine bones from
Laguna Niguel. It is possibly related to the Auklet, Aethia rossmoori Howard
1968, described from an ulna (LACM 18948) with referred distal end of
humerus (LACM 18949), collected in the Monterey Formation at Leisure
World in nearby Laguna Hills (locality LACM 1945). LACM 37686 resembles
the humerus of A. rossmoori in the rounded shaft, position of the brachial
impression with slight rise bordering it externally, and attachment of the an-
terior articular ligament facing more palmad than laterally. It is, however, 15%
larger than the specimen of A. rossmoori. Also, although the area of the tricipi-
tal grooves is abraded, the grooves appear to be less deeply incised than in
modern species of Aethia, or A. rossmoori.

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Contributions in Science

No. 290

Tribe Fraterculini
Genus and species indeterminate

Referred material. — Proximal end of humerus, LACM 42658, from lo-
cality LACM 7136, and distal end of humerus, LACM 37638, from locality
LACM 6902.

Discussion. — Both humeral fragments resemble this element in the puf-
fins (following Storer 1960:698, in segregating Cerorhinca in a tribe along with
Fratercula and Lunda). The proximal portion (LACM 42658) resembles the
type humerus of Cerorhinca minor Howard 1971 (LACM 15408), from the
Pliocene of Baja California, Mexico, in the presence of a ridge from the head to
the median crest, forming an internal border to the tricipital depression on the
anconal surface below the head, and in the extension of the median crest to the
border of the bicipital crest. In proximal breadth the Laguna Niguel specimen
is 12.0 mm, which is markedly larger than this dimension in C. minor (10.5
mm). It is, in fact, closer in size to a humerus (LACM Bi 696) of Lunda
cirrhata (Pallas 1769) which measures 12.2 mm in proximal breadth. Similarity
to Lunda is seen in the extension of the pectoral attachment to the deepest part
of the head, and, on the palmar side, the slight inset of the bicipital surface
border from that of the bicipital crest. Similarity to Fratercula corniculata
(Naumann 1821) is noted in the length of the bicipital surface, which becomes
slightly pointed at its distal extreme. The tricipital depression below the head is
shallower than in any of the specimens of Recent puffins at hand.

The distal portion of humerus (LACM 36738) also resembles this element
in the Fraterculini. Its size suggests that it may belong to the same species as
the proximal end discussed above.

The possiblity that these two bones might be assignable to Cerorhinca
dubia, described from the Late Miocene of Lompoc, was considered. How-
ever, their size seems to preclude this possibility. Miller (1925:116) gave the
measurements of length for the type leg bones of C. dubia as 60 mm (tibiotar-
sus) and 29 mm (tarsometatarsus). These measurements in a series of ten
specimens of C. monocerata (Pallas 1811) are 59.2 mm-60.0 mm, mean 62.9
mm (tibiotarsus), and 27.7 mm-30.6 mm, mean 29.1 mm (tarsometatarsus). The
humeri from Laguna Niguel are relatively larger in comparison with the same
series of skeletons of C. monocerata: proximal breadth (LACM 42658) 12.0
mm (C. monocerata 6.9 mm-11.3 mm, mean 10.6 mm); distal breadth (LACM
36738) 8.0 mm (C. monocerata 6.9 mm-7.8 mm, mean 7.2 mm).

Genus Praemancalla Howard 1976
Praemancalla wetmorei Howard 1976

No further material referable to this species has been found at Laguna
Niguel since the type description, which included: holotype humerus, LACM
42653, paratype ulna, LACM 32429, and referred proximal end of humerus,
LACM 32432, all from locality LACM 6906; complete radius, LACM 53907,

1978

Miocene Marine Birds

23

and scapular end of coracoid, LACM 37637, both from locality LACM 6902;
and proximal section of carpometacarpus, LACM 52216, from locality LACM
3185.

CONCLUSIONS

Seven families of marine birds are represented by the 39 identifiable avian
bones from the Late Miocene deposits in Laguna Niguel. In the following list
the numbers in parentheses indicate the number of specimens assigned to each
species.

Gaviidae — Loons

Gavia brodkorbi new sp. (1)

Diomedeidae — Albatrosses
Diomedea ? calif ornica (1)

Diomedea sp. indet. (2)

Procellariidae — Shearwaters
Puffinus barnesi new sp. (4)

Hydrobatidae — Storm Petrels
Oceanodroma sp. indet. (1)

Pseudodontornithidae — Extinct Bony-toothed Birds
Osteodontornis orri (3)

Sulidae — Boobies and Gannets
Morns lompocanus (7)

Morns magnns new sp. (4)

?Miosnla media (1)

Sulidae, spp. indet. (4)

Alcidae — Auklike Birds
?Uria sp. (1)

?Cepphns sp. (1)

?Aethia sp. (1)

Fraterculini, gen. and sp. indet. (2)

Praemancalla wetmorei (6)

Five of the above families (Diomedeidae, Procellariidae, Pseudodontor-
nithidae, Sulidae and Alcidae) are also represented in the larger collection of
Late Miocene birds recovered earlier from locality LACM 1945 in Leisure
World, Laguna Hills, about three miles north of Laguna Niguel (Howard
1968). However, few of the same species are listed from the two sites and none
of those described as new from locality LACM 1945 is found in the Laguna
Niguel area. From locality LACM 1945, 50% of the 120 bones identified are of
the Procellariidae (4 species), with Sulidae (3 species) and Alcidae (5 species)
constituting approximately 22% each. The remaining 6% include the Anatidae
(2 species), Diomedeidae (2 species) and ? Osteodontornis orri.

At Laguna Niguel only 10% of the 39 identified bones are procellariid (1

24

Contributions in Science

No. 290

species). The Sulidae are most abundant, making up 41%, followed by the
Alcidae 28%. Diomedea and Osteodontornis constituting 1Vi% each, are rela-
tively better represented than at locality LACM 1945. A loon (Gaviidae) and a
storm petrel (Hydrobatidae), not present at LACM 1945, complete the av-
ifauna. The Anatidae are not represented.

Although the deposits at both of these sites are in the Monterey Formation
of Late Miocene (Clarendonian correlative) age, comparison of the two av-
ifaunas strengthens the belief suggested by a comparison of the mancalline
auks (Howard 1976) that the fauna from Laguna Niguel represents a slightly
later time than that from locality LACM 1945. Not only is the mancalline auk,
Praemancalla wetmorei, from Laguna Niguel further specialized for wing-
propelled diving than is P. lagunensis from locality LACM 1945, but in the
Laguna Niguel sites there is less indication of persistence of Middle Miocene
species and a more definite representation of typical Late Miocene species. At
locality LACM 1945, Presbychen abcivus Wetmore 1930 and Puffinus priscus,
both described from the Middle Miocene Sharktooth Hill Bonebed, are listed.
Neither has been found at Laguna Niguel, although another species, Diomedea
calif ornica, described from the same locality, is tentatively identified. On the
other hand the typically Late Miocene Moms lompocanus, which is only tenta-
tively identified at locality LACM 1945, is the most abundant species at
Laguna Niguel. Osteodontornis orri, tentatively listed from LACM 1945, on
the basis of a single fragment, is definitely present at Laguna Niguel.

Some significance also may be attached to the occurrence of Gavia and
Oceanodroma at Laguna Niguel. This is the first Miocene west coast occurr-
ence of the loons (Gaviidae). The only previous record for Oceanodroma is O.
hubbsi Miller in the Capistrano Formation, which is now considered to be of
latest Miocene to early Pliocene (Hemphillian correlative) age (Fife 1974:19)
rather than Middle Miocene as interpreted by Miller (1951).

ACKNOWLEDGMENTS

Had it not been for the generosity of the North American Rockwell Land
Corporation (NARLAND), this study would not have been possible. Their
permission, even encouragement, to collect during their 1969 construction
project, and since then, to collect and quarry in adjacent hillsides on NAR-
LAND property, has resulted in amassing the present excellent representation
of Late Miocene fossils. I am also grateful to Mrs. Louise Hanson for granting
permission to collect on the Moulton Ranch adjacent to NARLAND property.

My thanks are extended to the following persons who have contributed
fossils for this study: Mr. and Mrs. Marion J. Bohreer and family, Mr. Paul R.
Kirkland, Mrs. Janet Price and family, Mr. W. Earl Calhoun, Mr. Rodney
Raschke, and Miss Jennifer Whistler.

I am grateful to Dr. Pierce Brodkorb of the University of Florida and Dr.
Storrs Olson of the United States National Museum of Natural History, for
consultation and for the loan of material for comparison; and to Dr. J. Howard

1978

Miocene Marine Birds

25

Hutchison, Museum of Paleontology, University of California, Berkeley for
the loan of comparative fossil material and permission to cast it. Others who
have contributed comparative material are: Dr. Everett C. Olson, University
of California at Los Angeles; Dr. Raymond A. Paynter, Jr., Museum of Com-
parative Zoology, Harvard University; Dr. Stuart Warter, California State
University, Long Beach; and the staff of the Museum of Vertebrate Zoology,
University of California, Berkeley.

I am continually indebted to the Natural History Museum of Los Angeles
County for the opportunity to work with the Museum’s collections, and to the
Paleontology staff, in particular, for their cooperation. In the present study.
Dr. Lawrence G. Barnes, who was in charge of some of the Museum’s excava-
tions at Laguna Niguel, has been a constant source of information and consul-
tation. Others to whom I am grateful for assistance in many ways are: Dr.
Theodore Downs, Dr. David Whistler, Robert McKenzie, andMichael Ham-
mer. The photographs were made by Lawrence Reynolds, Museum Photo-
grapher. The figures were prepared by Mary Butler, Graphic Artist at LACM.

The curation of this material was made possible under National Science
Loundation Grant DEB 7202014, to the Natural History Museum of Los
Angeles County Loundation.

LITERATURE CITED

Brodkorb, P. 1953. A review of the Pliocene loons. Condor, 55:211-214.

1963a. Miocene birds from the Hawthorne Formation. Quart. Jour. Florida

Acad. Sci., 26:159-167.

1963b. Catalogue of Fossil Birds, Part 1 (Archaeopterygiformes through Ar-

deiformes). Bull. Florida State Mus. Biol. Sci., 7:179-293.

1967. Catalogue of Fossil Birds, Part 3 (Ralliformes, Ichthyomithiformes,

Charadriiformes). Bull. Florida State Mus. Biol. Sci., 11:99-220.

Fife, D.L. 1974. Geology of the south half of the El Toro Quadrangle, Orange County,
California. California Div. Mines and Geol. Spec. Rep. 110:1-27.

Howard, H. 1957a. A gigantic “toothed” marine bird from the Miocene of California.
Santa Barbara Mus. Nat. Hist. Bull. Dept. Geol., 1:1-23.

1957b. A new species of passerine bird from the Miocene of California. Los

Angeles Co. Mus. Contrib. Sci. No. 9:1-16.

1958. Miocene sulids of Southern California. Los Angeles Co. Mus. Contrib.

Sci. No. 25:1-15.

1966a. A possible ancestor of the Lucas Auk (Family Mancallidae) from the

Tertiary of Orange County, California. Los Angeles Co. Mus. Contrib. Sci. No.
101:1-8.

1966b. Additional avian records from the Miocene of Sharktooth Hill, Califor-
nia. Los Angeles Co. Mus. Contrib. Sci. No. 114:1-11.

1968. Tertiary birds from Laguna Hills, Orange County, California. Los

Angeles Co. Mus. Contrib. Sci. No. 142:1-21.

1969. A new avian fossil from Kern County, California. Condor, 71:68-69.

26

Contributions in Science

No. 290

1971. Pliocene avian remains from Baja California. Los Angeles Co. Mus.

Contrib. Sci. No. 217:1-17.

1976. A new species of flightless auk from the Miocene of California (Al-

cidae:Mancallinae). Smithsonian Contrib., Paleobiol., 27:141-146.

Howard, H. and J.A. White. 1962. A second record of Osteodontornis, Miocene
“toothed” bird. Los Angeles Co. Mus. Contrib. Sci. No. 52:1-12.

Miller, L. 1925. Avian remains from the Miocene of Lompoc, California. Carnegie
Inst. Washington, Publ. No. 349:107-117.

1929. A new cormorant from the Miocene of California. Condor, 31:167-172.

1935. New bird horizons in California. Publ. Univ. California Los Angeles,

Biol. Sci. l(5):73-80.

1951. A Miocene petrel from California. Condor, 53:78-80.

1961. Birds from the Miocene of Sharktooth Hill, California. Condor, 63:399-

402.

1962. A new albatross from the Miocene of California. Condor, 64:471-472.

Milne-Edwards, A. 1874. Observations sur les oiseaux fossiles des faluns des Saucats et
de la mollasse de Leognan. Bib. Ecole Haute Etudes, Sect. Sci. Nat. XI(3): 1-12.

Shufeldt, R.W. 1896. Fossil bones of birds and mammals from Grotto Pietro Tamponi
and Grive-St. Alban. Proc. Acad. Nat. Sci. Phila. 1896, pp. 507-516.

Storer, R.W. 1956. The fossil loon, Colymboides minutus. Condor, 58:413-426.

1960. Evolution in the diving birds. Proc. XII Internat. Omith. Congress,

Helsinki, 1958, pp. 694-707.

Warter, S.L. 1976. A new osprey from the Miocene of California (Falconiformes:Pan-
dionidae). Smithsonian Contrib. Paleobiol. 27:133-139.

Wetmore, A. 1930. Fossil bird remains from the Temblor Formation near Bakersfield,
California. Proc. Calif. Acad. Sci., 4th Ser., 19(8):85-93.

1941. An unknown loon from the Miocene fossil beds of Maryland. Auk,

58:567.

1943. Fossil birds from the Tertiary deposits of Florida. Proc. New England

Zool. Club., 22:59-68.

Wilkinson, H.E. 1969. Description of an Upper Miocene albatross from Beaumaris,
Victoria, Australia, and review of fossil Diomedeidae. Mem. National Mus. Vic-
toria, 29:41-51.

Accepted for publication June 8, 1977.

50 7. 73
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MARCH 22, 1978

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Printed in the United States of America by Chapman's Phototypesetting on 70# Patina Book

THE NET-WINGED MIDGES OF EASTERN NORTH AMERICA,
WITH NOTES ON NEW TAXONOMIC CHARACTERS
IN THE FAMILY BLEPHARICERIDAE (DIPTERA)1

By Charles L. Hogue2

Abstract: The single genus Blepharicera Macquart of the family Blepharicer-
idae occurs in eastern North America, probably arriving historically via a continental
connection with western Europe, and is now distributed through Appalachia, New
England, the northern Great Lakes Region and southeastern Canada. Six species are
recognized, discussed and figured: B. capitata (Loew), cherokea new species, dim-
inutiva new species, similans (Johannsen), tenuipes (Walker), and williamsae (Alex-
ander). Six larval types also are known but none definitely can be associated with
adults (although three are provisionally identified). Pupal anatomy, with one excep-
tion, is homogeneous, defying species characterization.

The usefulness of taxonomic characters in the group is discussed including ap-
plication of several new terms.

INTRODUCTION

The family Blephariceridae is represented in eastern North America by the single
genus Blepharicera, in which six species are recognized presently. These species,
here designated as the “Tenuipes Group,” are extremely similar in all stages and
undoubtedly arose from a single ancestral form. Blepharicera is now known to occur
only in the northern hemisphere, ranging more or less continuously and narrowly
across Asia through northern Spain, the Alps and Balkans, Caucasus, Himalayas, to
Japan and North Korea. Species also are found disjunctly in northern Borneo, Thai-
land, Taiwan and neighboring China. This is a so-called “Tethyan distribution” and
suggests, within the tenets of plate tectonics, that the genus originated in Laurasia and
expanded to the south only as far as the northern margin of the Tethyan Sea.

Introduction to North America could have occurred via Europe-Newfoundland or
Bering connections. I favor the former route for the Tenuipes Group because of the
total lack of close relatives in western North America. (B. ostensackeni may form an
exception but it is not clearly a member of this group.)

The western species of Blepharicera could have evolved secondarily from a western
extension of this stock, but they are an assemblage of distinct species quite unlike the
eastern species and I think it much more likely that they arrived independently from

‘Review Committee for this Contribution
C. P. Alexander
Douglas Craig
Julian P. Donahue

2Senior Curator of Entomology, Natural History Museum of Los Angeles County, 900 Expo-
sition Boulevard, Los Angeles, California, 90007.

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one or more introductions over a Bering connection from eastern Asia. This thesis is
supported further by the parallel occurrence in western North America of the genera
Bibiocephala and Philorus , both of which have very close relatives in Japan and other
parts of Asia. Unfortunately, there is no information on the occurrence of the genus
in Alaska and northern and central Canada which could establish continuity between
the eastern and western forms.

The eastern Blepharicera are now known from as far west as eastern Minnesota
through southern Ontario and all the southeastern Canadian provinces, through New
England and down the Appalachian cordillera to its southern extreme in Georgia.
Included in this distribution are the Canadian provinces of Quebec, Ontario, New-
foundland, New Brunswick and Nova Scotia, and the American states of Maine,
Vermont, New Hampshire, Minnesota, New York, Massachusetts, Connecticut,
Pennsylvania, New Jersey, Maryland, Virginia, West Virginia, North Carolina, South
Carolina, Tennessee and Georgia. I predict that they may eventually be discovered
in extreme northeastern Alabama, eastern Kentucky and possibly Michigan and
Manitoba.

This largely congruent distribution and the extreme morphological similarity of all
the species has made it impossible, without rearings, to firmly establish associations
of the immature with the adult stages. As many as three species can occur at the same
locality, so that stages associated in single collections do not necessarily belong to
the same species. A further barrier to relating the larvae to the adults is the total
homogeneity of pupal morphology. Whereas identifiable pharate adults may be related
to pupae by dissection, there is no way to transcend the gap between larva and pupa.
The one exception to this is the distinctive form “F” with conspicuous median dorsal
convexities on the abdomen of larva and pupa. Unfortunately, the one mature form
“F” pupa available did not yield an identifiable female.

These difficulties have prevented me from definitely establishing stage associations
in any species, and all identifications of immatures cited are provisional. Associations
can be accomplished with certainty if specimens are individually reared, but this re-
quires special care, time, and equipment. Very probable associations can be estab-
lished if collecting is extensive and carefully done so that groups of lawae of all instars
present, and pupae in close proximity (on the same boulder or area of a large rock
face) and in the process of transformation, are preserved apart from other groups.
Unfortunately, the material available for this study was totally inadequate for this
application. In the taxonomic treatment below, I have relegated larvae to each of the
species very tentatively and state my reason, albeit tenuous, for each association. I
urge students of stream biology in the East to vigorously and carefully collect this
genus so that these vagaries can be removed.

TAXONOMIC CHARACTERS
Larval Chaetotaxy and Cuticular Sclerotizations

In my first attempts to separate the very similar larvae of the various species in this
complex, I became aware immediately of the need to utilize new characters. Previ-
ously, the chaetotaxy has been almost totally ignored in taxonomic studies of larval

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Blephariceridae Eastern North America

3

Blephariceridae, and no foundation existed upon which to assess possible useful char-
acter states. From the practical standpoint of need for a consistent terminology and
to reveal some points for species discrimination, I undertook a preliminary study of
the homologies (both serial and comparative between several genera) of the trunk
setae; study of the head setae has been deferred.

After examining the first instar larvae of A gathon comstocki (Kellogg) (Fig. 1 -west-
ern North America) and Neocurupira chiltoni (Campbell) (Fig. 2 - New Zealand) and
later instars of numerous genera, it became apparent that, like those of most nema-
tocerous families, blepharicerid larvae possess so-called primary sensilla which make
up the complete complement of the first instar larva and a much larger number of
secondary sensilla added in subsequent instars. The latter appear in addition to the
primary sensilla after the first molt and continue to proliferate until maturity when the
larva may be covered with a dense vestiture. In these later stages this homogeneity
and density obscures the recognition of the primary sensilla, but these often retain
some distinctive characteristic, such as larger alveolus, occurrence in pairs, darker
pigmentation, larger size, association with a tubercle or plate, or unique shape.

These patterns are very constant and follow a basic plan described as follows for
all except segments VIII - IX (refer to figures 1 - 2 and 30 for further explanation):
tergal sensilla (t) — most mesal series on the dorsum on all segments. Usually ob-
literated in later instars by proliferation.

subtergal (st) — next laterad to the tergal sensilla, on all segments. Always distin-
guishable in later instars by position, setiform shape and large alveolus.

tergopleural (tp) — two sensilla situated far laterad on all segments. Associated
with the dorsolateral tubercle or plate in those forms with these structures. A close-
set pair of long, setiform sensilla on the meso- and metathorax (although paired, not
serially homologous with abdominal “geminate setae”).

pre-dorsopseudopodal (pdpod) — in the first instar a single sensillum on the ex-
treme base of the pseudopod on the abdominal segments; an isolated seta on the
thorax. In later instars a group of setae on the anterobasal angle of the dorsal pseu-
dopod when this organ is present.

dorso-psetidopodal (“geminate setae”) (dpod) — a pair of conspicuous, always
large, setiform sensilla on the anterolateral angle of the trunk segments and lateral
margin of the cephalic division in instars II-IV; on the dorsal surface of the base of
the pseudopod in the first instar. These sensilla are located at the apices of the dorsal
pseudopods when this organ occurs and at the apex of the posterolateral lobe of the
anal division (which is a dorsal pseudopod anatomically).

pleural (p) — a single seta, variously positioned on the lateral margin. Usually com-
pletely obscured by the secondary setae in later instars.

sternopleural (sp) — on the thorax of Neocurupira chiltoni a single, heavy seta
between the conspicuous ventral substernal and lateral pleural hairs.

substernal (ss) — a large conspicuous seta on the venter of the thoracic segments,
laterad of the sucker and minute sternal (s) setae.

pedichetal (pd) — This group is contained within the ventral sucker and consists of
three minute setae. As shown by Craig (1967: fig. 13 and p. 202) the anteriormost
sucker belongs to abdominal segment I, the posteriormost to abdominal segment VI.

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No. 291

Pedichetal sensilla, therefore, appear to be wanting from the venters of the thorax and
anal division.

In addition to the relatively large setate sensilla, there are various minute sense
organs (intertergal-it, intersternal-is, intercalary -ic), some appearing as minuscule
hairlike processes, other as hairless rings (sensilla campaniforme). Their positions are
very constant in all instars and species, so they offer little more taxonomically than
reference points.

Segments VIII and IX of the anal division are recognizable dorsally by lines of
sensilla and sclerites. The complements of both, however, are reduced and the no-
menclature just given does not apply strictly. On segment VIII a tergal sensillum
remains distinct but the others are grouped into a single pleural group. On segment
IX, the terminal segment, only a pair of pleurals are apparent besides the conspicuous
terminal setae.

Both primary and secondary sensilla may take a variety of shapes, often charac-
teristic of genera and species. On the abdomen of the first instar of the two forms
which I have studied, all the primary sensilla are setiform except the tergal (t) and
the more lateral of the tergopleurals, both of which are lanceolate or coniform. This
is also true of the thorax except for the tergals and intertergals which are coniform
and the pleurals which are claviform.

This pattern persists in later larval instars of Blepharicera, except that the majority
of the dorsal secondary sensilla assume varied coniform, claviform or capitate shapes
and arrangements which confer to them diagnostic value.

The function of these organs is unknown. I doubt their sensory importance because
of their density. Their dorsal position and the frequent occurrence of specimens with
heavy growths of diatoms and algae and even mineral precipitates entangled and at-
tached to them, suggests a function similar to that of the “tectorial” setae and spines
of psychodid larvae (Vaillant 1959:41), be it to help anchor or hide the animal. The
roughened surface produced may also act physically to reduce resistance with the
water, like sclerotized projections which could have the same function as suggested
by Hora 1930:255. The condition would appear to be specialized and apomorphous
since few hairs is the rule in nematocerous and mecopterous larvae.

My study of chaetotaxy remains incomplete, requiring much material now unavail-
able of first instar larvae of many genera. I am continuing to work on an analysis of
the homologies of the sensilla and hope eventually to propose a system of nomencla-
ture useful to morphologists and taxonomists. For the present, I present detailed il-
lustrations of the patterns of sensilla in these few species with only general topographic
names for taxonomic reference.

Probably associated with these sensillar modifications is the tendency of many lar-
vae (although not Blepharicera) to form dorsal sclerotizations since the positions of
the latter generally coincide with the locations of the primary setae. The dorsal pseu-
dopod (as in Agathon and Philorus) for example, always bears the large paired dorso-
pseudopodal (dpod) setae at its apex. The blepharicerid taxonomist is advised, how-
ever, to homologize such plates, tubercles and other sclerotized processes with great
caution because these structures take similar forms in unrelated species and genera.

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Blephariceridae Eastern North America

5

Pupal Structures

With the exception of Species F, I have been unable to discover characters on the
pupa which permit species discrimination. Part of the problem is the lack of complete
material of all species. This most conservative of life stages shows no divergence of
structure, even in detail, among these very closely related Eastern species. Size varies
continuously from the smallest, diminutiva, to the largest, williamsae, so cannot be
used as a discrete objective character.

Adult Structures

Aside from the number of bristles on the parietal sclerite of the head, which varies
from none in most males to up to 50 in females of williamsae, there is little in the
general anatomy of either sex to reliably distinguish the species.

It should be emphasized that the latter statement is also largely true of size and
external body coloration, although these characters formed the basis of Alexander’s
key (1963:52-53). After trying to reconcile the several hundred adults used in the
present study with this key, I have come to the conclusion that coloration in this
species group is a character of very limited use because, (1) it varies continuously
from species to species (smaller species tending to be paler than larger) and (2) its
interpretation is too easily confused by the condition and age of the specimen when
killed, presence of grease, and the angle of lighting and viewing. Regarding this last
point, I should explain that the integument induces interference phenomena in light
rays reflected from it and it displays spectral colors when viewed from oblique angles.
Thus, the pigmentary colors are obscured and distorted by other than perpendicular
viewing and the thorax seems to change in color from gray to brown as the specimen
is rotated under the stereoscopic microscope.

The most useful species characters in this group, as with most nematocerous Dip-
tera, are to be found in the genitalia. In the male the following three character com-
plexes are those applied mainly: (1) shape of the apex and shaft of the parameres;
(2) shape of the apices of the penis filaments; and (3) shape of the IXth tergite lobes.
All tend to be simpler in the smaller species, a fact which throws some light on the
phylogeny of the group by suggesting that size differences may have generated al-
lometric structures that present reproductive barriers (see REMARKS under diminutiva).

Genitalic features of the females are less well marked than in the males. B. similans
is highly distinctive in lacking a normal median spermatheca (correlated with the
overly long and apically modified median penis filament in the male) but the number
of these organs remains three in all the other forms. Of use primarily are the following
characters although even these all seem to be subject to such variation as to make
identification of isolated or atypical specimens often impossible: (1) number and dis-
tribution of macrochaetae on the VUIth sternite lobes, (2) shape of the median fold,
especially the median basal sclerotization thereof, (3) sclerotizations of the spermathe-
cal ducts.

An additional fairly definitive and constant character is the shape of an organ which

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I interpret from gross morphology as the accessory gland. Unfortunately it is some-
times difficult to discern in slide mounts, but when evident its outline takes a variety
of consistent forms.

PROCEDURE

All statistics given are means with the range in parentheses. Unless otherwise noted,
the sample size for each set of measurements was normally 10 for wings and legs,
2-5 for head structures. Wherever the sample was less than 10 (e.g. diminutiva),
ranges are not given in the descriptions. Because of the small series available from
specific localities, it was usually necessary to take data from mixed populations. Body
length of larvae was measured only on prepupal individuals (pupal branchiae visible).
Measurements in text and on the figures are in millimeters.

Proportions of leg segments are given as “progressive proportions” meaning that
each segment is taken relative to that proximal rather than to one standard segment.
This makes comparisons easier and more accurate since the segments are progressively
shorter to a small degree or nearly equal in length; greatly differing and disjunct
lengths need not be compared to one another.

In the case of antennal segment proportions, the basal flagellar segment is used as
the standard since it is more conveniently and accurately measured than either the
scape or pedicel which are both globular and difficult to orient consistently.

Material is listed separately for the stages. That for the larvae includes pupae which
accompanied the specimens when collected but which were insufficiently mature to
make an adult determination. Pupae containing pharate adults are treated as adult
records and recorded in their section of the species accounts.

Some new morphological terms have been introduced. They appear in bold-face
when first used in the text and their application explained in the following figures:
female genitalia (Fig. 27); head capsule (Figs. 14-15); larva (Figs. 1-2, 30).

TAXONOMY-ADULTS
Blepharicera, Tenuipes Group

DIAGNOSIS

Coloration: General. — Integument well sclerotized, generally gray-brown or
deep red-brown with dull gray (plumbeous) overtones. Membranes and appendages
pale yellow to medium brown. All major sclerite surfaces opalescent; head sclerite
surfaces also finely pruinose. Head. — Face, occiput and basal portion of clypeus gray,
pruinose. Suprafrontal carina and distal portion of clypeus dull orange to brown.
Callis oculi shiny black. Antenna. — Scape brownish yellow, pedicel and flagellum
dark brown. Mouthparts generally orange to yellow, palpus grading to brown apicad.
Thorax. — Anterior pronotum, mesoscutum and scutellum gray to gray brown; pos-
terior pronotum, posterolateral comers of scutum usually, and prescutellar area of

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Blephariceridae Eastern North America

7

scutum often, contrasting pale brown or yellow. No other well-defined lines or pat-
terns on scutum. Preepistemum, epistemum and meron similar to scutum but lighter
and sometimes with reddish-brown tints; remainder of pleuron light brown to yellow.
Legs. — Brownish-yellow basad, grading to dark brown distad. Wing. — Hyaline in
both sexes; veins brown. Haltere. — Stem yellow or light brown basad; knob dark
brown. Abdomen. — Tergites and stemites similar to scutum. Pleural membranes yel-
low to brown. Genitalia yellow orange to dark brown.

Size: Generally medium to small Blephariceridae. Wing length, range 3.2 - 9.5

mm.

Head: Structure. — General structure as usually found in family. Antennal fossae
approximate, mesofrons very narrow.

Female. Clypeus length/width 2. 0-2. 2. Suprafrons narrow, suprafrontal carina
strongly convex. Parietal sclerite broad (0.7 times width of infrafrons), trapezoidal
in shape. Ocellar lobes joined to form a sessile tubercle; lenses inserted on lobes
apically. Eyes. — Approximate at level immediately anterior to ocelli, interocular dis-
tance equal to diameter of one ommatidium; upper division well differentiated from
lower; callis oculi broad, anterior portion striate. Size of upper division about equal
to lower in area, 13-16 rows of ommatidia along maximum dorsal arc of upper di-
vision; upper ommatidia 1.4-2. 5 times diameter of lower. Proboscis. — Short, about
0.7 times head width. Mandibles present.

Male. Clypeus length/width ratio slightly greater than female, 2.4-3. 6. Supra-
frons broad, slightly convex mesally. Parietal sclerite broad, but narrower than in fe-
male (0.2-0. 4 times width of infrafrons), rectangular in shape. Ocellar lobes distinct;
lenses inserted on lobes apically. Eyes. — Disjunct at level immediately anterior to
ocelli, ipterocular distance equal to diameters of 3-5 upper ommatidia; upper division
well differentiated from lower; callis oculi absent except for narrow anterior portion.
Size of upper division one-fourth to one-half of lower in area, 8-16 rows of ommatidia
as in female. Proboscis. — Short, about 0.6 times head width. Mandibles absent. Pal-
pal segments five, proportions constant, segment 1 small and fused at base to rostrum,
3 and 4 about equal and each slightly longer than 2, 5 three to four times length of
2, about equal to 2-3 combined. Chaetotaxy . — Similar in sexes, except parietal bris-
tles usually absent in male. Major setal groups present as follows: clypeal, variably
sparse over most of surface; parietal, few to numerous bristles; vertical, few short
bristles; postvertical, numerous short bristles over dorsal arc of occipital foramen;
occipital, numerous long bristles; postgenal, numerous long bristles. Ocular bristles
short.

Antenna. Structure. — Moderately long, about 1.5 times head width, 15-seg-
mented. Scape subspherical, pedicel conopiform, flagellomeres elongate-cylindriform
in female, stouter and compressed in male. Chaetotaxy . — Scape with group of nu-
merous long slender bristles mesoventrally. Pedicel with small dorsal and ventral
groups of short bristles. Flagellomeres with dense vestiture of macrotrichia (these
more numerous on male) and scattered placoid sensilla on apical segments (confined
to apex of terminal segment in male).

Thorax: Chaetotaxy. — Setal patterns incompletely studied. Scutellar bristles
dense laterally, incomplete or absent mesally. Wing. — -Venation typical for genus.

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Dorsal macrotrichia on veins R5 and M3 complete, M1+2 and M4 apically only. Legs.
— Segments simple, tarsal segments 5 all unmodified. Tibial spurs absent from fore
and mid legs, 1 or 2 on hind leg. Claws simple, similar and not enlarged.

Female Genitalia: VUIth sternite lobes broadly rounded, with or without setae.
Oviscapt subquadrate, base slightly wider than apex, inner piece poorly developed.
Spermathecae three (rarely two) in number, generally ovoid in shape with no or poorly
developed necks.

Male Genitalia: Segments VIII and IX not specially modified; IXth tergite
lobes prominent, subquadrate in form, posterior margin with a small acute projection.
Basistyle short and broad, slightly longer than outer dististyle. Latter a simple, elon-
gate lobe, slightly depressed on inner surface apically. Inner dististyle a narrow curved
spatulate lobe. Phallosome vesica small, spherical; apodeme a simple, vertical flange.
Parameres and penis filaments varied in form.

KEY TO SPECIES
Adult Males

1. Median penis filament distinctly longer than laterals and with asymmetrical,
hooked apex. Wing length 4.1 mm (3.4-4.65). Apex of paramere an incurved,
asymmetrically bifurcate finger. IXth tergite lobe shape as in figure 22.

similans

Median penis filament about same length as laterals, apex unmodified or only
slightly enlarged. Apex of paramere and IXth tergite lobe varied in shape but not
as above. Wing length 3. 2-6. 6 mm 2

2. Posterior margin of tegmen deeply incised on either side of midline. Apex of par-
amere a dorso-mesally directed hook. IXth tergite lobe as in figures 24 or 26.

Wing length 5.4 mm (4. 4-6. 2) tenuipes

Posterior margin of tegmen entire. Other characters varied but not as above ... 3

3. Inner margin of IXth tergite lobe greatly expanded mesad (posteromesal comer

obliterated). Larger species, wing length 4.8 mm or greater 4

Inner margin of IXth tergite lobe straight from posteromesal comer to base, not
greatly expanded (posteromesal comer angulate). Smaller species, wing length 4.8
or less 5

4. Inner wall of apex of paramere an incurved, acute process extending well beyond
aperture. IXth tergite lobe shape as in fig. 28. Wing length 5.6 mm (4. 9-6.6)

williamsae

Inner wall of apex of paramere very slightly produced, truncate; aperture terminal.
IXth tergite lobe shape as in figure 16. Wing length 5.1 mm (4. 8-5. 3)
capitata

5. Posterior margin of IXth tergite lobe with submesal projection, general shape as

in figure 18. Wing length 4.4 mm (4. 0-4. 8) cherokea

Posterior margin of IXth tergite lobe straight, oblique, general shape as in figure
20. Wing length 3.8 mm (definitive range undetermined) diminutiva

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Blephariceridae Eastern North America

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Adult Females

1 . Two spermathecae, median reduced to a cylindrical rod. Wing length 6.0 mm (5.4-

7.3) similans

Three spermathecae. Wing length varied 2

2. Ducts of lateral spermathecae sclerotized for a short length before genital aperture.

Wing length 5.95 mm (5. 2-6. 4) capitata

Ducts of spermathecae membranous throughout. Wing length varied 3

3. A large species, wing length 8.1 mm (6.75-9.5). Thoracic scutum with a con-
trasting light brown quadrate area anterior to scutellum. VUIth stemite lobe of

genitalia devoid of setae williamsae

Smaller species, wing length 7.1 mm or less. Thoracic scutum generally unicol-
orous. VUIth stemite lobe of genitalia usually with at least one major seta ... 4

4. VUIth sternite lobes contiguous, median fold V-shaped. Very small species, wing

length 5.1 mm (definitive range undetermined) diminutiva

VUIth sternite lobes disjunct, median fold U-shaped. Larger species, wing length
usually greater than 6.0 mm (5.5-7. 1) 5

5. Mesoscutum reddish-brown. Accessory gland a straight tube with irregular anterior

sclerotizations. Wing length 6.0 mm (5. 5-6. 5) cherokea

Mesoscutum dull gray (plumbeous). Accessory gland triangular, very broad pos-
teriorly, membranous anteriorly; wing length 6.6 mm (6.0-7. 1) tenuipes

Blepharicera capitata (Loew)

Blepharoptera capitata Loew 1863:298-299. LECTOTYPE by present designation, female,
District of Columbia, Washington, 1866, C. R. Osten Sacken, (MCZ, No. 16124).

Blepharocera capitata , Osten Sacken 1895:161. Kellogg 1903 (partim).

Blepharocera tenuipes, Aldrich 1905:172 (original synonymy; attributed to Osten Sacken). Kel-
logg 1907:12. Alexander 1963:56.

Blepharicera capitata, Stone 1965:99.

Blepharocera separata Alexander 1963:54-55. NEW SYNONYMY.

DESCRIPTIONS
Adult female (Figs. 9, 17)

Size: A medium-sized Blepharicera', measurements as follows: Wing length 5.95

(5. 2-6. 4), width 1.90 (1.60-2.10). Head width 1.06. Labrum length 0.31. Palpal seg-
ments 2-5, lengths: 0.10, 0.13, 0.145, 0.33. Leg segment lengths as below:

fore

mid

hind

femur

3.6 (3.2-3.95)

3.6 (3.15-4.0)

4.85 (4.25-5.4)

tibia

3.1 (2.75-3.35)

3.05 (2. 3-3. 3)

4.45 (3.95-4.9)

tarsus 1

1.41 (1.25-1.64)

1.46 (1.30-1.60)

1.78 (1.64-1.91)

2

0.79 (0.68-0.88)

0.78 (0.66-0.85)

0.64 (0.58-0.71)

3

0.52 (0.44-0.55)

0.53 (0.48-0.56)

0.39 (0.34-0.43)

4

0.30 (0.28-0.34)

0.31 (0.25-0.39)

0.26 (0.20-0.30)

5

0.30 (0.25-0.34)

0.31 (0.25-0.36)

0.29 (0.24-0.33)

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Thoracic Coloration: Anterior pronotum, mesoscutum and scutellum reddish-
brown with slight gray overtone especially mesad; preepisternum, episternum and
meron like mesoscutum but lighter; remainder of pleuron pale yellow.

Head (Fig. 9): Parietal sclerite with few (1-8) setae, restricted to lower half of
sclerite. Distal four palpal segment proportions: 1.0, 1.4, 1.1, 2.3. Antennal segment
proportions: 0.87, 0.87, 1.0, 0.70, 0.79, 0.77, 0.75, 0.75, 0.77, 0.78, 0.76, 0.76,
0.73, 0.74, 1.31; ultimate segment 1.70 length of penultimate.

Legs: Tibial spur formula 0-0-2 (lesser very small). Progressive proportions of
leg segment lengths as below:

foreleg: 1.00, 0.86, 0.46, 0.56, 0.66, 0.58, 1.01
midleg: 1.00, 0.85, 0.48, 0.53, 0.67, 0.59, 1.01
hindleg: 1.00, 0.92, 0.40, 0.36, 0.61, 0.68, 1.09

Genitalia (Fig. 17): VUIth stemite lobes broadly rounded, median fold U-
shaped, very shallow; lobe with 0-4 (usually 1) macro setae, in posterolateral region.
Spermathecae 3 in number, equal in size and oval in shape; necks absent; ducts var-
iably sclerotized, usually with posterior-most sections of two lateral ducts narrowed,
smoothly sclerotized and pigmented. Accessory gland broad, anterior portion sharply
expanded spherically or semi-rectangularly.

Adult male (Figs. 3, 16)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 5. 1 (4. 8-5.3),
width 1.68 (1.60-1.80). Head width 0.86 (0.82-0.89). Labrum length 0.22 (0.20-
0.24). Palpal segments 2-5, lengths: 0.10, 0.13, 0.14, 0.38. Leg segment lengths as
below:

femur
tibia
tarsus 1
2

3

4

5

fore

3.1 (2. 8-3.4)
2.8 (2.65-3.05)
1.48 (1.38-1.63)
0.78 (0.69-0.88)
0.54 (0.60-0.48)
0.29 (0.26-0.31)
0.27 (0.25-0.31)

mid

3.6 (3.1-5.85)
2.8 (2.65-3.00)
1.47 (1.31-1.61)
0.73 (0.64-0.82)
0.52 (0.45-0.58)
0.29 (0.25-0.31)
0.28 (0.23-0.30)

hind

4.35 (3. 9-4.7)
4.0 (3.65-4.4)
1.53 (1.31-1.75)
0.52 (0.44-0.61)
0.34 (0.25-0.38)
0.24 (0.20-0.29)
0.25 (0.21-0.26)

Head (Fig. 3): Parietal sclerite without setae. Distal four palpal segment pro-
portions: 1.0, 1.4, 1.6, 4.2. Antennal segment proportions: 0.70, 0.71, 1.0, 0.97,
1.02, 1.05, 0.95, 0.98, 0.96, 1.0, 0.94, 0.94, 0.88, 0.85, 1.06; ultimate segment 1.2
length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segments as
below:

foreleg: 1.00, 0.91, 0.52, 0.53, 0.70, 0.53, 0.94
midleg: 1.00, 0.78, 0.52, 0.50, 0.70, 0.55, 0.96
hindleg: 1.00, 0.92, 0.38, 0.34, 0.64, 0.72, 1.02

Genitalia (Fig. 16): IXth tergite lobes prominent, parallel; lobe shape irregular,
outer comer a rounded right angle, posterior margin projecting beyond outer angle

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Blephariceridae Eastern North America

11

as a broadly rounded lobe which gradually curves into inner margin to base of lobe;
bristles numerous and regular over entire dorsal surface. Paramere broad basally, grad-
ually tapering apically and curving outwardly; apex truncate (inner margin slightly
produced); aperture terminal. Penis filaments 3 in number, simple truncate rods, me-
dian slightly longer and stouter than laterals and with small apical collar or cap, all
shorter than parameres.

VARIATION

Some females fail to exhibit the typical pigmented, sclerotized sections of the
posterior extremes of both of the lateral spermathecal ducts. Neither duct may be so
modified, but in any case this region of the duct is always narrowed and smooth
walled in contrast to the wholly membranous tubules found in other species.

Males occur in which the median penis filament is scarcely stouter than the lat-
erals, and the inner margin of the IXth tergite lobe is expanded. This character state
conforms to that of the holotype of Blepharicera separata Alexander. Since these
specimens are identical in all other ways with the presumed male of capitata, I choose
to interpret these differences as minor and therefore synonymize Alexander’s form.

REMARKS

Some authors have synonymized capitata with tenuipes, but apparently without
basing their decision on a thorough examination of the types of capitata. I have as-
sembled five specimens which appear to be from the original series (collected by Osten
Sacken) including two females labelled “type” in the collection of MCZ. No males
are among these although indicated in Loew’s description. Believing Loew to be in
error in citing the sex of his material, I have selected one of the latter two females
as a lectotype.

Superficially these females are indistinguishable from typical tenuipes. However,
the genitalia are distinct and define a separate species to which I assign equally distinct
males found at the type locality where no other species are known to occur.

DISTRIBUTION (Fig. 37)

This species is found along the entire Appalachian chain and beyond to the north
into New Brunswick. The form named separata by Alexander may appear anywhere
in this range and apparently is no more than an occasional variant.

MATERIAL

Adults: DISTRICT OF COLUMBIA. Washington, 1866, C. R. Osten Sacken (1 9 LEC-
TOTYPE: MCZ No. 16124; 4 9 PARALECTOTYPES: MCZ No. 16124, AMNH, USNM).
GEORGIA, Rabun County. Pine Mountain, 1400 feet, 15 May 1957, W. R. M. Mason (21 6 ,
6 9: CNC). Towns County. Hiawassee, 16 June 1945, P. W. Fattig (1 6: USNM). MARY-
LAND, Cecil County. Octorora Canyon, 17 June 1939, E. G. Fisher (1 9 :ANSP). Prince
George’s County. Beltsville, 28 May 1916, W. L. McAtee (2 9:USNM). NEW HAMPSHIRE,
Grafton County. Franconia, Mrs. Slosson (1 cLUSNM). PENNSYLVANIA, Monroe County.
Delaware Water Gap, 15 June 1934, C. P. Alexander (1 cUUSNM). VIRGINIA, Fairfax
County. Great Falls, 21 June 1931, A. L. Melander (1 9 :USNM); Trammel’s Landing, Potomac
River, [above Great Falls, now Riverbend County Park] 26, 28 May 1935, A. Stone (1 6 , 2
9 iUSNM). VERMONT, Chittenden County. Smuggler’s Notch, 18 June 1927, C. P. Alexander
(9 :USNM).

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Atypical {separata form). — MAINE, Somerset County. Bingham, along the Kennebec
River, 17 July 1937, C. P. Alexander (3 6 PARATYPES: CPA). NEW BRUNSWICK, North-
umberland County. Boiestown, 13 July 1931, J. M. Aldrich (lcCUSNM). NORTH CARO-
LINA, Madison County. Hot Springs (IcCAMNH). Swain County. Soco Valley, Cherokee, 20
June 1954, J. W. Green (1$:CAS). NEW YORK, Fulton County. Sport Island, Sacandaga
River, 18 June 1914, C. P. Alexander (29 :USNM). PENNSYLVANIA, Lebanon County. Ono,
7 June 1940, A. L. Melander (1 $ :USNM). VERMONT, Windham County. Dummerston, 14
July 1908 (19 :USNM).

Blepharicera cherokea new species

DIAGNOSIS

Adults

Blepharicera cherokea is well defined only by characters found in the male gen-
italia, principally the shapes of the IXth tergite lobes and apex of the paramere. The
former resembles that of tenuipes , but the small median lobe on the hind margin is
decidedly more mesal and the outer corner less pronounced. The latter presents a
complex appearance in tenuipes, being a dorso-mesally directed, asymmetrical hook,
formed by elaboration of the inner wall, while in cherokea both inner and outer walls
terminate almost equally, giving the structure an oblique, truncate apex.

Female specimens with the anterior portion of the accessory gland irregularly
sclerotized are tentatively assigned to this species. There is little else to separate them
from tenuipes, the most closely related entity.

ETYMOLOGY

This species is named in honor of the Cherokee Indian Nation, in whose home-
land it resides.

DESCRIPTIONS

Adult female (Figs. 10, 19)

Size: A moderately- small Blepharicera', measurements as follows: wing length
6.0 (5. 5-6. 5), width 1.96 (1.85-2.15). Head width 1.10. Labrum length 0.32. Palpal
segments 2-5, lengths: 0.10, 0.14, 0.15, 0.49. Leg segment lengths as below:

fore

mid

hind

femur

3.7 (3.35-4.1)

3.6 (3.15-4.0)

4.8 (4.35-5.1)

tibia

3.1 (2. 7-3. 4)

2.9 (2.55-3.05)

4.4 (3. 9-4.6)

tarsus 1

1.44 (1.31-1.53)

1.48 (1.24-1.61)

1.90 (1.63-2.06)

2

0.75 (0.63-0.85)

0.72 (0.60-0.81)

0.66 (0.63-0.75)

3

0.46 (0.38-0.55)

0.45 (0.38-0.54)

0.38 (0.29-0.43)

4

0.28 (0.23-0.34)

0.27 (0.23-0.29)

0.26 (0.23-0.28)

5

0.29 (0.25-0.35)

0.29 (0.25-0.33)

0.29 (0.26-0.31)

Thoracic Coloration: Anterior pronotum, mesoscutum and scutellum reddish-
brown with gray overtone, especially mesally; lateral corners of anterior and posterior

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pronota yellow; preepistemum, epistemum and meron like mesoscutum but lighter;
remainder of pleuron pale yellow.

Head (Fig. 10): Parietal sclerite with numerous (20-30) setae over entire median
portion. Distal four palpal segment proportions: 1.0, 1.4, 1.5, 4.9. Antennal segment
proportions: 0.85, 0.85, 1.0? 0.69, 0.77, 0.85, 0.77, 0.77, 0.75, 0.77, 0.69, 0.69,
0.62, 0.69, 1.08; ultimate segment 1.56 length of penultimate.

Legs: Tibial spur formula 0-0-2, (lesser spur of hind leg about one-half the size
of greater). Progressive proportions of leg segments as below:
foreleg: 1.00, 0.82, 0.47, 0.52, 0.61, 0.61, 1.04
midleg: 1.00, 0.82, 0.51, 0.49, 0.63, 0.60, 1.07
hindleg: 1.00, 0.92, 0.43, 0.35, 0.58, 0.68, 1.12

Genitalia (Fig. 19): VUIth sternite lobes wide-set, median fold shallowly U-
shaped; each lobe with few to numerous setae. Spermathecae 3 in number, equal in
size and ovoid in shape; necks short. Accessory gland a straight tube with irregular
anterior sclerotizations.

Adult male (Figs. 4, 18)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 4.4 (4. 0-4. 8),
width 1.48 (1.35-1.70). Head width 0.67. Labrum length 0.19. Palpal segments 2-5,
lengths 0.10, 0.14, 0.14, 0.39. Leg segment lengths as below:

fore

femur 2.8 (2.55-3.1)

tibia 2.6 (2.35-2.9)

tarsus 1 1.46 (1.33-1.63)

2 0.73 (0.64-0.81)

3 0.51 (0.44-0.58)

4 0.25 (0.23-0.28)

5 0.23 (0.20-0.25)

mid

2.9 (2.6-3.55)

2.5 (2.25-2.8)
1.39 (1.26-1.50)
0.69 (0.60-0.78)
0.49 (0.41-0.54)
0.25 (0.21-0.26)
0.23 (0.20-0.25)

hind

3.75 (3. 4-4.3)

3.5 (3. 2-3. 9)
1.35 (1.23-1.48)
0.45 (0.40-0.50)
0.28 (0.25-0.31)
0.18 (0.16-0.23)
0.20 (0.18-0.25)

Head (Fig. 4): Parietal sclerite without setae. Distal four palpal segment pro-
portions: 1.0, 1.4, 1.4, 4.0. Antennal segment proportions: 0.63, 0.63, 1.0, 0.94,
1.16, 1.18, 1.10, 1.10, 1.10, 1.18, 1.18, 1.18, 1.10, 1.18, 1.26; ultimate segment

1 .06 length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segments as
below:

foreleg: 1.00, 0.92, 0.50, 0.50, 0.70, 0.49, 0.92
midleg: 1.00, 0.86, 0.55, 0.50, 0.70, 0.50, 0.94
hindleg: 1.00, 0.93, 0.39, 0.33, 0.62, 0.65, 1.13

Genitalia (Fig. 18): IXth tergite lobes prominent, parallel; lobe shape subquad-
rate, outer posterior corner a rounded right angle, not prominent; pointed lobe pro-
jecting from hind margin displaced mesad; bristles numerous and regular over entire
dorsal surface. Tegmen entire, with a slight apico-median carina. Paramere broad
basally, tapering regularly and gradually to apex; apex simple, obliquely truncate
(inner wall only very slightly longer than outer); aperture terminal. Penis filaments

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3 in number, equal in length, shorter than parameres, all simple truncate rods, the
median slightly stouter than laterals.

VARIATION

The numer of setae on the VUIth stemite lobe of the female varies from 3 to 8.
This indicates possible heterogeneity in the small sample available to me. Associated
material is definitely needed to confirm the identity of the females of this species.

REMARKS

Association of the sexes is in some doubt because no females are available from
the type locality. Collecting from populations containing known males should easily
resolve the question.

DISTRIBUTION (Fig. 39)

This species is known only from the type locality and collections near the south-
ern end of the Appalachians (Georgia and North Carolina).

MATERIAL

Types. — HOLOTYPE 6 (genitalia on slide no. CLH 75-166; head on slide no. CLH 75-
239: NORTH CAROLINA, Macon County. Highlands, Clear Creek, 3200 feet, 1-2 July 1958,
J. G. Franclemont. 25 PARATYPE 6 : same data as holotype. All deposited at CU.

Additional specimens. GEORGIA, Dawson County. Amicalola Falls No. 5, 14 June 1960,
G. W. Byers (19$ :KU). NORTH CAROLINA, Swain County. Bryson City, 3-5000 feet, May
1936, R. C. Shannon (lcUUSNM); Cherokee, Soco Valley, 20 June 1954, J. W. Green
(1 $ CAS); Cherokee, 2000 feet, 24 May 1957, W. R. M. Mason (1$:CNC). TENNESSEE,
Sevier County. Greenbrier Cove, 2000 feet, Great Smoky Mountains National Park, 18 May
1957, J. R. Vockeroth (2 <3:CNC).

Blepharicera diminutiva NEW SPECIES

DIAGNOSIS

Adults

Although this is the smallest species of Blepharicera it is set apart from the other
members of its group primarily by unique features of the male genitalia; a simple,
subquadrate IXth tergite lobe rather than the complex and marginally expanded shapes
of all the other species, and the paramere apex with merely a thin, truncate extension
of the shaft in place of being forked or spined in various ways. The females are
difficult to distinguish because the most reliable character (shape of the accessory
gland) is often obscure. This structure is an indefinite, membranous tube in this species
whereas in the other species it assumes a variety of forms as a result of sclerotization.
Also characteristic, but ill-defined, are the close-set VUIth tergite lobes, meeting with
a V-shaped median fold between; in all the other species the median fold is much
broader, U-shaped and often with angular basal corners.

ETYMOLOGY

The name is derived from the Latin adjective diminutivus , meaning tiny, in ref-
erence to the species’ small size.

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Blephariceridae Eastern North America

15

DESCRIPTIONS

Adult female (Figs. 11, 21)

Size: A very small Blepharicera\ measurements as follows: Wing length 5.1,
width 1.63. Head width 0.97. Labrum length 0.32. Palpal segments 2-5, lengths:
0.09, 0.12, 0.13, 0.44. Leg segment lengths as below:

fore

mid

hind

femur

3.3

3.1

4.2

tibia

2.7

2.55

3.9

tarsus 1

1.25

1.2

1.6

2

0.595

0.545

0.52

3

0.39

0.34

0.33

4

0.215

0.215

0.21

5

0.23

0.23

0.22

Thoracic Coloration: Anterior pronotum and mesoscutum generally brown-
ish-gray; scutellum brownish-yellow; corner of posterior pronotum yellow; preepi-
stemum, epistemum and meron like mesoscutum but lighter; remainder of pleuron
brownish-yellow.

Head (Fig. 1 1): Parietal sclerite with numerous setae over entire median portion.
Distal four palpal segment proportions: 1.0, 1.3, 1.4, 4.9. Antennal segment pro-
portions: 0.85, 0.77, 1.0, 0.85, 0.92, 0.77, 0.77, 0.77, 0.77, 0.69, 0.69, 0.69, 0.54,
0.54, 1.0; ultimate segment 1.85 length of penultimate.

Legs: Tibial spur formula 0-0-2 (lesser spur of hind leg minute). Progressive
proportions of leg segment lengths as below:

foreleg: 1.00, 0.82, 0.47, 0.48, 0.66, 0.55, 1.07
midleg: 1.00, 0.82, 0.46, 0.46, 0.62, 0.63, 1.07
hindleg: 1.00, 0.92, 0.42, 0.32, 0.64, 0.64, 1.05

Genitalia (Fig. 21): VUIth sternite lobes close-set, median fold V-shaped; each
lobe with a few (4-6) macro setae. Spermathecae 3 in number, equal in size and ovoid
in shape; necks short to nearly absent; ducts membranous throughout. Accessory gland
a simple membranous tube, without definite shape.

Adult male (Figs. 5, 20)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 3.8, width
1 .29. Head width 0.65. Labrum length 0. 17. Palpal segments 2-5, lengths: 0.09, 0.09,
0.10, 0.35. Leg segment lengths as below:

fore

mid

hind

femur

2.3

2.4

3.2

tibia

2.1

2.05

2.95

tarsus 1

1.13

1.07

1.17

2

0.51

0.50

0.35

3

0.35

0.35

0.22

4

0.19

0.19

0.145

5

0.17

0.18

0.17

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Head (Fig. 5): Parietal sclerite without setae. Distal four palpal segment pro-
portions: 1.0, 1.0, 1.1, 3.9. Antennal segment proportions: 0.90, 0.70, 1.0, 0.80,
0.90, 1.0, 0.90, 0.90, 0.80, 0.90, 0.80, 0.90, 0.80, 0.90, 1.10; ultimate segment 1.2
length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segment lengths
as below:

foreleg: 1.00, 0.91, 0.54, 0.45, 0.69, 0.54, 0.89

midleg: 1.00, 0.85, 0.52, 0.46, 0.70, 0.54, 0.94

hindleg: 1.00, 0.91, 0.40, 0.30, 0.63, 0.66, 1.15

Genitalia (Fig. 20): IXth tergite lobes prominent, slightly divergent; lobe shape
subquadrate, a slight lobe projecting from midposterior margin, inner margin straight;
bristles numerous and regular over entire dorsal surface. Tegmen entire. Paramere
with a broad base, straight and gradually tapering shaft; apex a short, truncate, thin
extension, with a slightly produced inner margin; aperture terminal. Penis filaments
3 in number, equal in length, shorter than parameres; all simple, similar, capitate
rods.

REMARKS

The simplicity of several characters in both sexes (shape of the male IXth tergite
lobe, V-shaped female median fold) of this species, its overall smallness and its rarity
raise the possibility that it may actually constitute the negative allomorphic variant
of another, sympatric form, perhaps cherokea. However, until this can be positively
demonstrated, I think the entity should be recognized as a species.

DISTRIBUTION (Fig. 38)

This species has the most restricted range of the eastern B lepharicera. It is known
only from localities clustered in the southernmost sector of the Blue Ridge Mountains
located at the junction of the states of Georgia, South Carolina and North Carolina.

MATERIAL

Types. — HOLOTYPE 8 (genitalia on slide no. CLH 75-177; wing on slide no. CLH 75-
243): NORTH CAROLINA, Macon County. Highlands, Clear Creek, 3200 feet, 1 July 1958,
J. G. Franc lemont; ALLOTYPE 9 (genitalia on slide no. CLH 75-155; mid and hind legs on
slide no. CLH 75-248); 1 PARATYPE 8 and 1 PARATYPE 9 : same data as holotype. All
deposited at CU.

Additional specimens. — NORTH CAROLINA, Transylvania County. Lake Toxaway, 12
July 1957, J. G. Chillcott (1 cLCNC). GEORGIA, Rabun County. Addie Branch, east fork of
Chattooga River, 2400 feet, 1 August 1957, J. G. Chillcott (1 9 :CNC); Rabun Bald, 3000 feet,
14 July 1957, J. G. Chillcott (19 :CNC); Tallulah Falls, 10 June 1910 (1 9 :CU); Satalah, 2000
feet, 1 July 1957, J. R. Vockeroth (2<?:CNC).

Blepharicera similans (Johannsen)

Blepharocera similans Johannsen 1929: 123-124. Type locality, Massachusetts, Hampshire
County, Orient Springs, near Amherst. Holotype male, CU.

Blepharocera similans , Johannsen 1934:50. Alexander 1963:55.

Blepharicera similans, Stone 1965:99.

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Blephariceridae Eastern North America

17

DESCRIPTIONS

Adult female (Figs. 12, 23)

Size: A medium-sized Blepharicera\ measurements as follows: Wing length 6.0
(5. 4-7. 3), width 2.05 (1.69-2.75). Head width 1.08. Labrum length 0.37. Palpal seg-
ments 2-5, lengths: 0.11, 0.14, 0.14, 0.45. Leg segment lengths as below:

fore

mid

hind

femur

3.5 (2.3-4.25)

3.76 (3.25-6.65)

4.8 (4.25-5.65)

tibia

3.0 (2.75-3.55)

2.9 (2.6-3.25)

4.3 (3.7-5.05)

tarsus 1

1.44 (1.26-1.68)

1.30 (1.15-1.44)

1.70 (1.41-2.0)

2

0.71 (0.57-0.81)

0.67 (0.54-0.76)

0.60 (0.46-0.73)

3

0.41 (0.26-0.50)

0.41 (0.30-0.46)

0.36 (0.28-0.41)

4

0.30 (0.29-0.33)

0.26 (0.21-0.30)

0.26 (0.21-0.30)

5

0.34 (0.28-0.43)

0.33 (0.30-0.40)

0.34 (0.31-0.39)

Thoracic Coloration: Anterior pronotum and mesoscutum generally dark
gray-brown; scutellum distinctly lighter than mesoscutum, brown, darker laterally;
preepistemum, epistemum and meron reddish brown; remainder of pleuron light-
brown.

Head (Fig. 12): Parietal sclerite with numerous setae (15-20) on lateral half of
surface. Distal four palpal segment proportions: 1.0, 1.25, 1.2, 4.0. Antennal segment
proportions: 0.60, 0.80, 1.0, 0.78, 0.71, 0.75, 0.66, 0.63, 0.63, 0.60, 0.59, 0.59,
0.59, 0.54, 0.93; ultimate segment 1.72 length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segment lengths
as below:

foreleg: 1.00, 0.86, 0.47, 0.49, 0.58, 0.75, 1.12
midleg: 1.00, 0.77, 0.44, 0.51, 0.61, 0.63, 1.28
hindleg: 1.00, 0.89, 0.40, 0.35, 0.60, 0.72, 1.32

Genitalia (Fig. 23): VUIth stemite lobes broadly rounded, median fold deep,
U-shaped, basal comers angular; lobe without setae. Spermathecae 2 in number, the
median one reduced to a vestigial, sclerotized bar; laterals ovoid, equal in size and
in shape; necks very short; ducts membranous throughout. Accessory gland cylindrical
with a slightly inflated median zone.

Adult male (Figs. 6, 22)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 4. 1 (3.4-4.65),
width 1.32 (1.20-1.50). Head width 0.73 (0.69-0.82). Labrum length 0.21 (0.18-
0.24). Palpal segments 2-5, lengths 0.08, 0.12, 0.12, 0.42. Leg segment lengths as
below:

fore

mid

hind

femur

2.6 (2.2-3.05)

2.7 (2.2-3. 2)

3.55 (2.9-4.25)

tibia

2.4 (1. 9-2.9)

2.3 (2. 0-2. 7)

3.3 (2.7-3.85)

tarsus 1

1.31 (0.85-1.56)

1.26 (0.98-1.48)

1.23 (0.98-1.45)

2

0.62 (0.44-0.75)

0.60 (0.49-0.74)

0.39 (0.33-0.48)

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3

4

5

fore

0.42 (0.28-0.61)
0.22 (0.13-0.28)
0.22 (0.16-0.28)

mid

0.41 (0.33-0.48)
0.22 (0.16-0.28)
0.22 (0.16-0.28)

hind

0.23 (0.16-0.30)
0.16 (0.13-0.19)
0.20 (0.13-0.24)

Head (Fig. 6): Parietal sclerite without setae. Distal four palpal segment pro-
portions: 1.0, 1.3, 1.4, 4.1. Antennal segment proportions: 0.77, 0.68, 1.0, 0.79,
0.82, 0.86, 0.79, 0.76, 0.72, 0.74, 0.75, 0.81, 0.80, 0.83, 1.11; ultimate segment
1.3 length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segments as
below:

foreleg: 1.00, 0.93, 0.55, 0.47, 0.68, 0.51, 1.02
midleg: 1.00, 0.86, 0.54, 0.47, 0.68, 0.54, 1.00
hindleg: 1.00, 0.94, 0.37, 0.32, 0.59, 0.70, 1.25

Genitalia (Fig. 22): IXth tergite lobes prominent, parallel; lobe shape irregular,
outer comer a rounded right angle, posterior margin projecting posteriorly as an acute
prominence, inner margin evenly and shallowly curved to the slightly constricted base;
bristles numerous and regular over entire dorsal surface. Tegmen entire. Paramere
broad basally, tapering gradually to two-thirds, then abruptly narrowing ectally to
point where ental wall terminates, apex extended beyond aperture as a gradually in-
curved, asymmetrically bifurcate finger. Penis filaments 3 in number, all shorter than
paramere but median heavier and longer than laterals and with a hooked apex.

VARIATION

No notable variations were observed in this species, which is the most constant
and distinctive member of the Tenuipes Group.

REMARKS

The female of this species is unique in the genus in the absence of the median
spermatheca. The organ is replaced by an elongate structure apparently modified es-
pecially to receive the male’s extra long, hooked, median penis filament. The func-
tional significance of this adaptation is unknown.

DISTRIBUTION (Fig. 40)

There are collections of similans from localities along the entire Appalachian
chain from Maine to South Carolina and Tennessee. The species is sympatric with
all the other members of its group, but would appear to be the least likely one to
hybridize because of the genitalic peculiarities mentioned above. This also, perhaps,
explains its lack of variability.

MATERIAL

Adults. — CANADA. NOVA SCOTIA, Colchester County. Truro, 4 July 1913, R. Matheson
(3 6 :CU). QUEBEC, Gatineau County. Wakefield, 9 July 1946, G. E. Shewell (2 <3:CNC);
18 September 1928, 25 July 1926, F.'p. Ide (1 6, 1 9:ROM); 21 July 1926, G. S. Walley
(1 6, 1 9:CNC); 1 July 1959, J. R. Vockeroth (1 6, 1 9:CNC). Huntingdon County. Covey
Hill, 8 July 1924, G. S. Walley (1 cLCNC). Laprarie County. Laprarie, 8 July 1924, G. S.
Walley (1 cLCNC). UNITED STATES. GEORGIA, Dawson County. Amicalola Falls No. 5,

1978

Blephariceridae Eastern North America

19

14 June 1960, G. W. Byers (1 9:KU). Rabun County. Satalah, 2000 feet, 1 July 1957, J. R.
Vockeroth (5 8 :CNC). Warwoman Creek, 1500 feet, 31 July 1957, J. G. Chillcott (2 8,
1 9 :CNC). MASSACHUSETTS, Hampshire County. Amherst, 25 June 1927 (1 8 :BMNH),
Orient Springs, 375 feet, 30 May 1926, C. P. Alexander (1 9 :CPA). NEW HAMPSHIRE, Coos
County. Gorham, 18 July 1929, G. S. Walley (5 8 :CNC). Moose Creek, Gorham, 19 July 1929,
J. McDunnough (1 9 :CNC). Dolly Copp, White Mountains, 29 August 1937, A. L. Melander
(1 8 , 5 9 :USNM). County Unknown. White Mountains, Morrison (28 :USNM). NEW YORK,
Essex County. Wilmington, Adirondacks, 31 July 1929, A. L. Melander (4 8, 6 9 :USNM).
High Falls, Ausable River, 5 July 1938. C. P. Alexander (2 cLUSNM). Lake Placid, 28 July
1929, A. L. Melander (5 c?:USNM); 1 July 1922, J. M. Aldrich (3 c?:USNM); 19 July 1962,
J. R. Vockeroth (3 9 :CNC). NORTH CAROLINA, A very County. Linville, 3200 feet, 21 June
1939, C. P. Alexander (3 9 :CPA). Macon County. Clear Creek, Highlands, 3200 feet, 1-2 July
1958, J. G. Franclemont (18 <J:CU). Wayah Bald, 3500 feet, 6 July 1957, J. R. Vockeroth
(1 9 :CNC). Wayah Gap, 3500 feet, 10 August 1957, J. G. Chillcott (2 9 :CNC). Swain County.
Smokemount, 2 July 1941, A. L. Melander (1 cLUSNM). Bryson City, 3-5000 feet, May 1936,
R. C. Shannon (1 8 :USNM). Wilkes County. Stone Mountain State Park, 10 July 1973, G. K.
Pratt (2 8,2 9:HDP). TENNESSEE, Sevier County. Chimneys, Great Smoky Mountains Na-
tional Park, 8 July 1941, A. L. Melander (1 8 :USNM). Smoky Mountains, 23 October 1938,
I, Williams (1 9:CPA). VIRGINIA, Page County. Luray, 24 June 1933, A. L. Melander
(2 8 :USNM).

Pupae (pharate adults dissected). — UNITED STATES. NEW YORK, Jefferson County.
Watertown, 30 November 1881, J. Q. Adams, No. 2213 (8 9,3 pupae, 20 larvae:USNM).
Ulster County. Vemooy Kill, 3/4 mi NW Wawarsing, 28 July 1974, P. & B. Wygodzinsky (26
pupae, 8 larvae:AMNH). NORTH CAROLINA, Swain County. Noland Creek, Great Smoky
Mountains National Park, 16 August 1973, 24 May 1974, R. L. Green (1 pupa :LACM). TEN-
NESSEE, Sevier County. Greenbrier Cove, Smoky Mountains National Park, 22 May, 10 Sep-
tember (?) 1938, A. C. Cole (34 pupae:UT). VIRGINIA, Giles County. Stoney Creek, 1800,
2000 feet, 26 May 1962, J. R. Vockeroth (18 pupae, 48 larvae:CNC). County Unknown. Dark
Hollow Falls, Shenandoah National Park, 30 July 1949, E. A. Chapin (15 pupae:USNM).

Asindulum tenuipes Walker 1848:86. Type locality, CANADA, Ontario, Albany River, holotype
female, BMNH.

Blepharocera capitata, Kellogg 1900:1903 (partim). Johannsen 1903:333-336.

Blepharocera tenuipes, Comstock & Comstock 1895:432-436. Aldrich 1905:172. Kellogg
1907:12. Johannsen 1934:54. Alexander 1963:56.

Blepharicera tenuipes. Stone 1965:99.

DESCRIPTIONS
Adult female (Figs. 14-15, 27)

Size: A medium-sized Blepharicera', measurements as follows: Wing length 6.6
(6.0-7. 1), width 2.1 (1.78-2.4). Head width 1.08 (1.01-1.44). Labrum length 0.38.
Palpal segments 2-5, lengths: 0.11, 0.15, 0.17, 0.46. Leg segment lengths as below:

Blepharicera tenuipes (Walker)

fore

mid

hind

femur
tibia
tarsus 1

3.9 (3. 4-4.3)
3.3 (3. 0-3. 6)
1.57 (1.30-1.79)
0.82 (0.68-0.94)
0.52 (0.45-0.60)
0.32 (0.24-0.37)
0.39 (0.33-0.46)

3.8 (3.35-4.2)
3.2 (2.9-3.45)
1.31 (0.89-1.65)
0.77 (0.56-0.94)
0.49 (0.41-0.59)
0.33 (0.27-0.40)
0.40 (0.33-0.46)

5.4 (4.9-5. 8)

4.9 (4.3-5. 3)

2.10 (1.84-2.21)
0.75 (0.68-0.84)
0.44 (0.40-0.49)
0.33 (0.28-0.37)
0.36 (0.28-0.43)

2

3

4

5

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Thoracic Coloration: Anterior pronotum and mesoscutum generally dull gray
(plumbeous), comer of posterior pronotum and scutellum contrasting light brown,
latter darker laterad; preepistemum, epistemum and meron similar to scutum but paler
and with reddish-brown cast; remainder of pleuron brownish-yellow.

Head (Figs. 14-15): Parietal sclerite with numerous setae (22-32) over entire
median surface. Distal four palpal segment proportions: 1.0, 1.4, 1.1, 2.7. Antennal
segment proportions: 0.92, 1.00, 1.0, 0.85, 0.85, 0.85, 0.77, 0.77, 0.81, 0.85, 0.77,
0.77, 0.69, 0.62, 0.92; ultimate segment 1.50 length of penultimate.

Legs: Tibial spur formula 0-0-2 (spurs subequal). Progressive proportions of leg
segment lengths as below:

foreleg: 1.00, 0.84, 0.47, 0.52, 0.63, 0.62, 1.22
midleg: 1.00, 0.85, 0.40, 0.59, 0.64, 0.67, 1.21
hindleg: 1.00, 0.90, 0.43, 0.36, 0.59, 0.75, 1.09

Genitalia (Fig. 27): VUIth stemite lobes broadly rounded, median fold shallow
and wide at base; lobe with a few (usually 6-7) macro setae. Spermathecae 3 in num-
ber, equal in size and ovoid in shape; necks very short; ducts membranous throughout.
Accessory gland triangular, very broad posteriorly, tapering strongly anteriorly.

Adult male (Figs. 7, 24-26)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 5.4 (4. 4-6. 2),
width 1.83 (1.56-2.16). Head width 0.86 (0.81-0.94). Labrum length 0.25 (0.22-
0.29). Palpal segments 2-5, lengths 0.12, 0.15, 0.16, 0.44. Leg segment lengths as
below:

fore

mid

hind

femur

3.3 (2. 9-3.7)

3.5 (3. 0-3. 9)

4.7

(4. 1-5.0)

tibia

3.1 (2. 7-3.4)

2.95 (2. 6-3. 3)

4.3

(3. 8-4.6)

tarsus 1

1.60 (1.24-1.85)

1.53 (1.28-1.76)

1.68

(1.38-1.87)

2

0.82 (0.71-0.92)

0.77 (0.67-0.89)

0.56

(0.46-0.68)

3

0.55 (0.39-0.70)

0.53 (0.41-0.63)

0.38

(0.33-0.57)

4

0.31 (0.27-0.35)

0.31 (0.26-0.35)

0.25

(0.21-0.35)

5

0.31 (0.24-0.41)

0.29 (0.26-0.33)

0.27

(0.23-0.33)

Head (Fig. 7): Parietal sclerite without setae. Distal four palpal segment pro-
portions: 1.0, 1.25, 1.1, 2.75. Antennal segment proportions: 0.63, 0.63, 1.0, 0.69,
0.81, 0.81, 0.78, 0.81, 0.81, 0.81, 0.78, 0.80, 0.78, 0.76, 0.91; ultimate segment
1.2 length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segments as
below:

foreleg: 1.00, 0.92, 0.52, 0.51, 0.67, 0.56, 1.00
midleg: 1.00, 0.85, 0.52, 0.50, 0.69, 0.58, 0.94
hindleg: 1.00, 0.92, 0.39, 0.33, 0.68, 0.66, 1.08

Genitalia (Figs. 24-26): DCth tergite lobes prominent, parallel; lobe shape ir-
regular, generally quadrate, outer corner pronounced, rounded, posterior margin hor-
izontal, with a mesally directed lobe near the middle; inner corner right angulate, the
margin continuing straight to base; bristles numerous and regular over entire dorsal

1978

Blephariceridae Eastern North America

21

surface. Tegmen deeply emarginate on either side of strong, median, vertical carina.
Paramere broad basally, only slightly tapering apically and nearly straight; both inner
and outer walls complete to apex; apex complex, a dorso-mesally directed, asym-
metrical hook; aperture subterminal. Penis filaments 3 in number, simple truncate
rods, of equal diameter, all shorter than paramere.

VARIATION

The southernmost record for typical tenuipes is Stoney Creek, Giles County,
Virginia. All specimens from localities further south exhibit certain states approaching
those found in cherokea. The most common involves the shape of the IXth tergite lobe
of the male genitalia in which the small mesal projection of the posterior margin is dis-
placed strongly mesad (Fig. 24) as in cherokea (Fig. 18). In other specimens, the
posterolateral comer of this same structure is abnormally pronounced, a condition also
reminiscent of cherokea or williamsae (Figs. 18, 28). The tegmen in some of these
same specimens has very shallow excisions on either side of the postero-median car-
ina. In all the other species the posterior margin of the tegmen is entire and without
a carina except in cherokea. All variants preserve the typical tenuipes form of the
apex of the paramere although there are a few cherokea specimens in which the nor-
mally simple apex is slightly extended and contorted in a fashion similar to the com-
plexities found here in tenuipes. The existence of these intermediates suggests close
genetic ties between cherokea and tenuipes. Although the possibility of hybridization
exists present material is inadequate to reveal the real nature of these relationships.

Variations in the female genitalia are less well defined and so far involve only
three specimens in which the number of macro setae on the VUIth stemite lobes and
configuration of the median fold are unusual.

REMARKS

This is the most common species of Blepharicera in Eastern North America.
Since it is the oldest named regional form, the other similar species, especially cap-
itata, have been confused with it. These two are indistinguishable externally, and the
genitalic characters cited in the key must be used to separate them.

DISTRIBUTION (Fig. 41)

A map plot of the records below shows tenuipes to be the most widespread spe-
cies in the group. It is mainly northern, although atypical specimens, tentatively as-
signed to this taxon, appear through the southernmost portions of the range of the
genus in Tennessee, North Carolina and Georgia (see VARIATION above).

There are no records over a large area north of the Great Lakes, but widely
disjunct collections, to the west at the Albany River, Ontario and eastern Minnesota,
and abundant suitable habitats are cause to believe that the Great Lakes region is also
populated with tenuipes. It is the only species found in the vicinity of Ithaca, New
York where it has long been well known to entomologists and students at Cornell
University.

MATERIAL

Adults.— CANADA. NOVA SCOTIA, Kings County. Kentville, 13 July 1924, R. P.
Gorham (1 9 :CNC). Victoria County. Baddeck, 27 June, 1 July 1936, J. McDunnough (3

22

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No. 291

9 :CNC). ONTARIO, Carleton County. Ottawa, 19 July 1924, G. S. Walley (1 9 :CNC); 22-
23 June 1943, G. S. Walley (4 9 :CN C); 24 June 1943, A. Brooks (9 9:CNC); 15 June 1951,
J. F. McAlpine (1 8, 5 9:CNC); 13 July 1956, J. R. Vockeroth (13 9:CNC). Shirley Bay,
Ottawa, 23 June 1961, J. A. Downes (4 8, 4 9:CNC); Brittania, Ottawa, 15 June 1938, G.
E. Shewell (1 <3 :CNC). Remic Rapids, Ottawa River, 18-21 June 1949, G. E. Shewell (2
cTCNC). Wentworth County. Dundas, 29 May 1960, Taylor and Wood (1 8, 1 9 :MMU).
QUEBEC, Brome County. Glensutton, 17 June 1936, G. S. Walley (1 9 :CNC). Knowlton, 8
July 1929, L. J. Milne (1 9 :CNC); 13 June 1930, G. S. Walley (1 <J:CNC). Gatineau County.
Aylmer, 10 June 1924, 26 June 1938, C. H. Curran (33 9 :AMNH). Chelsea, 20-25 June 1916
(2 9 :CU). Hull, 26 June 1920, 8 July 1923, Miss. Cramp, C. H. Curran (1 6, 1 9:CNC). Old
Chelsea, 22 July 1956, J. R. Vockeroth (3 9:CNC). Royal Ottawa Golf Club, Ottawa, 14, 31
July 1924, 27 June 1932, F. P. Ide, G. S. Walley (5 9:CNC, ROM). Wakefield, 21 August
1925, 21, 26 July 1926, G. S. Walley (1 8, 3 9:CNC); 14 June 1926, 28 July 1926, 4 June
1935, 18 September 1928, 9 July 1946, F. P. Ide (12 8, 5 9:CNC, ROM); 1 July 1959, J. R.
Vockeroth (1 8 :CNC). Huntingdon County. Covey Hill, 11, 28 June, 4 July 1924, 15, 18 June
1927, G. S. Walley (2 8,2 9 :CNC). Kamouraska County. Parke Reserve, 9 July 1957, G. E.
Shewell (1 8 :CNC); 8 August 1957, W. R. M. Mason (2 9 CNC). Laprarie County. Laprarie,

8 July 1924, G. S. Walley (8 9:CNC). Rouville County. Richelieu, 5 July 1927, G. S. Walley
(1 9:CNC). Stanstead County. Mount Orford, 14 July 1936, G. E. Shewell (1 9 :CNC).
UNITED STATES . CONNECTICUT , Fairfield County . Redding, 11 June 1929, A. L. Melander
(1 9 :USNM). MAINE, Hancock County. Bar Harbor, 4 June 1913, C. W. Johnson (1 9:USNM).
MINNESOTA, Cook County. Minnesota “F. S.”, Hovland, 23 June 1968, E. F. Cook, N.J.
mosquito trap (1 cUUSNM). NEW JERSEY, Morris County. Brookside, July, A. J. Weidt (3

9 :AMNH, USNM). NEW HAMPSHIRE, Coos County. Bretton Woods, 1 July 1936, A. L.
Melander (2 cUUSNM); Gorham, 18, 20 July 1929, j. McDunnough (2 9 :CNC); Jefferson
(black light trap), 10 July 1964 (1 9:UNH). Grafton County. Benton, 6 July 1931, A. L. Me-
lander (5 9 :USNM); Franconia, A. T. Slosson (1 8 ,2 9 :USNM, AMNH); Noxon Camp, 2000
feet, North Haverhill, 7 July 1931, J. M. Aldrich (3 9 :USNM). County unknown. White Moun-
tains, July (1 8 :USNM). NEW YORK, Erie County. Colden, 9 July 1922, 23 July 1916, M.
C. Van Duzee (2 9 CAS). Essex County. Lake Placid, 1 July 1922 (5 8 :USNM); 28 Julyl929,
A. L. Melander (1 d:USNM). Wilmington Notch, Adirondacks, 2 July, J. M. Aldrich (2
9 :USNM). Fulton County. Sacandaga, 15 June 1928, C.P. Alexander (2 8 , 1 9 :USNM). Lewis
County. Gomer Hill, 21 June 1963, W. W. Wirth (21 8, 3 9 :USNM). Singing Waters Picnic
Area, 21 June 1963, W. W. Wirth (3 8, 1 9 :USNM). Jefferson County. Watertown, J. Q.
Adams (1 9 :USNM). Saint Lawrence County. Raquette River, Colton, 24 June 1963, W. W.
Wirth, (1 cUUSNM). Tompkins County. Coy Glen, 18 June 1894, 18 June 1895, 12 June 1920,
J. M. Aldrich (part) (48,4 9:LACM, USNM, CU). Fall Creek, Ithaca, 10-11 June 1920 (8
9 :CU). Ithaca: 23 June 1885, O. E. Pearce (1 9:CU); 24 June 1901, J. M. Aldrich (21
9 :USNM, AMNH); 21 June-7 July 1901 (2 <3, 8 9 :BMNH). 12, 27-28 June 1901 (20 9:UCR,
CU); 14 June 1907 (1 9:CU); July 1908 (4 9 :ANSP); 31 May 1914 (13 8, 3 9:USNM); 14
June 1914, 12-13, 17, 24 June 1915 (6 9 :CU); 1, 10, 17, 21, 25 June 1916 (3 8, 17 9 :KU,
USNM, PSU, KS, CU); 13 June 1917, O. A. Johannsen (8 9 :PSU, CU); 14 June 1920 (1
9 :CU); 10 June 1925 (1 9 :CU); 3 June 1967, R. Silberglied (1 9 :CU); 1-7 July, N. Banks
(1 9 :OS); (3 9:USNM); May (6 <?:USNM); no dates: (1 8, 29: USNM, CU); (1 9 :CU);
Forest Home, Ithaca, 30 May 1940, A. Stone (3 <3:USNM). Taghanic, Ithaca, 26 June 1920,
M. D. Leonard (3 9:CU). PENNSYLVANIA, Adams County. Arendtsville, 27 May, 10 June
1920, S. W. Frost (3 9 :CU). County unknown. Castle Rock, 19 June 1910 (1 9 :USNM). VIR-
GINIA, Fairfax County. Great Falls, 19 June 1910, 24 May 1914, 23 May 1918, 21 June 1931,
30 May 1939, F. K. Knab, W. L. McAtee, A. L. Melander, C. T. Greene (7 8, 14 9 :USNM).
Giles County. No. 5, Mountain Lake Biological Station, 1800 feet, 18 June 1965, G. W. Byers
(2 9 :KU). Stoney Creek, 2000 feet, 26 May 1962, J. G. Chillcott (9 8,5 9 :CNC). VER-
MONT, Addison County. East Middlebury, 14 June 1929, Bishopp No. 8435 (1 8 :USNM).
Bennington County. Peru, 15 July 1931, A. L. Melander (1 9 :USNM). Windham County. Hal-
ifax Gorge, 8 June 1973, H. D. Pratt (11 <3:HDP, CPA). LOCALITY UNKNOWN. (1
9 :INHS).

1978

Blephariceridae Eastern North America

23

Atypical adults. — GEORGIA, Walker County. Cloudland Canyon State Park, 8 May 1952,
G. S. Walley et al. (20 6, 7 $:CNC). NORTH CAROLINA, Macon County. Highlands, 3-
5000 feet, May 1936, R. C. Shannon (3 cLUSNM). Same locality, 3200 feet, 2 July 1958, J.
G. Franclemont (2 9 :CU). Swain County. Deep Creek at Deep Creek Campground, Great Smoky
Mountains National Park, Bryson City, 21 May 1970, Wiggins and Yamamoto, 700365 (2
<3:ROM). TENNESSEE, Sevier County. Great Smoky Mountains National Park, 18 May 1957,
J. R. Vockeroth (1 <3:CNC); Greenbrier Cove, 2000 feet, 22 May 1938, I. Williams (1
<3:USNM). VIRGINIA, Bath County. Blowing Springs Camp, 8 mi W Warm Springs, 18-20
May 1963, Field and Flint (2 c?, 1 9:USNM). Page County. Luray, 21-24 June 1933, R. L.
Melander (2 <?, 3 9:USNM).

Pupae (pharate adults dissected). — CANADA. ONTARIO, Wentworth County. Spencer
Creek, 27 May 1960, D. M. Wood (1 6, 6 9, 25 pupae, 4 larvae:CNC). QUEBEC, Brome
County. Sutton Mountain Creek, 11 July 1929, G. S. Walley (16 pupae:CNC). UNITED
STATES. MAINE, Somerset County. Pleasant Pond Stream, Caratunk, 24 June 1966, R. & D.
Koss (21 pupae:JH). MASSACHUSETTS, Worcester County. Fitchburg, 27 May 1871 (44 pu-
pae:USNM). NEW YORK, Lewis County. Gomer Hill, 21 June 1963, W. W. Wirth (19 6, 44
pupae, 15 larvae:USNM).

Pupae (atypical pharate adults dissected). — UNITED STATES. MARYLAND, Frederick
County. Little Catoctin Creek, 1 mi N Harmony, 30 May 1958, P. H. Freytag (2 pupae, 1
larva:OSU). Little Hunting Creek, 4, 17, 25 May 1958, P. H. Freytag (5 pupae, 24 larvae:OSU).
Stream 1/2 mi E Yellow Springs, 31 May 1958, P. H. Freytag (8 pupae:OSU). NORTH CAR-
OLINA, Swain County. Deep Creek Campground, Great Smoky Mountains National Park, Bry-
son City, 21 May 1970, Wiggins & Yamamoto #700365 (4 pupae, 2 larvae:ROM). Noland
Creek, Great Smoky Mountains National Park, 24 May 1974, R. L. Green (1 pupa:LACM).
VIRGINIA, Giles County. Stoney Creek, 1800, 2000 feet, 25 May 1962, J. R. Vockeroth (33
pupae, 1 larva:CNC). Madison County. White Oak Creek, Shenandoah National Park, 24 June
1959, B. D. Burks (5 pupae, 4 larvae:USNM).

BlEPHARICERA WILL1AMSAE ALEXANDER

Blepharocera williamsae Alexander 1953:43, Type locality Tennessee, Sevier County, above
Greenbrier Cove, Mount Leconte, Great Smoky Mountains, 4200 feet elevation. Holotype
female and allotype male in Alexander collection.

Blepharocera williamsae , Alexander 1963:57. Redescription.

Blepharicera williamsae. Stone 1965:99.

DESCRIPTIONS
Adult female (Figs. 13, 29)

Size: A medium-sized Blepharicera: measurements as follows: Wing length 8.1
(6.75-9.5), width 2.57 (2. 1-2.9). Head width 1.27. Labrum length 0.48. Palpal seg-
ments 2-5, lengths: 0.16, 0.19, 0.18, 0.58. Leg segment lengths as below:

fore

mid

hind

femur

4.2 (4. 1-5.4)

4.45 (3.85-5.1)

6.2

(5.4-7. 1)

tibia

3.5 (3.3-4.55)

3.6 (3.1-4.15)

5.5

(4.85-6.1)

tarsus 1

1.84 (1.56-2.09)

1.61 (1.36-1.80)

2.25

(1.96-2.46)

2

0.89 (0.81-1.08)

0.81 (0.65-0.99)

0.77

(0.66-0.88)

3

0.55 (0.41-0.64)

0.50 (0.38-0.59)

0.47

(0.40-0.54)

4

0.36 (0.30-0.44)

0.34 (0.30-0.41)

0.33

(0.28-0.39)

5

0.46 (0.40-0.60)

0.43 (0.35-0.58)

0.40

(0.34-0.50')

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No. 291

Thoracic Coloration: Anterior pronotum and mesoscutum generally reddish-
brown with slight gray overtone especially mesad; comer of posterior pronotum, pos-
teromesal quadrate area of scutum and scutellum markedly lighter than mesoscutum,
pale yellowish-brown; outer comers of scutellum darker; remainder of pleuron brown-
ish-yellow.

Head (Fig. 13): Parietal sclerite with numerous (35-50) setae over entire median
surface of sclerite. Distal four palpal segment proportions: 1.0, 1.2, 1.1, 3.6. Antennal
segment proportions: 0.77, 0.78, 1.0, 0.77, 0.80, 0.80, 0.80, 0.80, 0.80, 0.80, 0.74,
0.74, 0.68, 0.62, 0.99; ultimate segment 1.60 length of penultimate.

Legs: Tibial spur formula 0-0-2 (spurs subequal). Progressive proportions of leg
segment lengths as below:

foreleg: 1.00, 0.82, 0.53, 0.48, 0.62, 0.66, 1.26
midleg: 1.00, 0.81, 0.45, 0.50, 0.62, 0.67, 1.27
hindleg: 1.00, 0.89, 0.33, 0.34, 0.61, 0.63, 1.18

Genitalia (Fig. 29): VUIth stemite lobes disjunct, broadly rounded, median
fold U-shaped, the basal comers angular; lobe without macro setae, a pair of micro
setae on either side of midline on bridge between lobes. Spermathecae 3 in number,
equal in size and ovoid in shape; necks very short; ducts membranous throughout.
Accessory gland a slightly inflated, elongate tube.

Adult male (Figs. 8, 28)

General: Character states as given for female except as follows:

Size: Smaller than female; measurements as follows: Wing length 5.6 (4. 9-6. 6),
width 1.85 (1.65-2.05). Head width 0.85. Labrum length 0.25. Palpal segments 2-5,
lengths: 0.10, 0.14, 0.14, 0.52. Leg segment lengths as below:

fore

mid

hind

femur

3.4

(2.75-4.0)

3.7

(3.45-4.2)

4.8

(4. 5-5. 5)

tibia

3.3

(3. 1-3.7)

3.1

(2.85-3.55)

4.5

(4. 1-5.3)

tarsus 1

1.78

(1.60-2.09)

1.61

(1.45-1.89)

1.70

(1.48-2.00)

2

0.83

(0.68-0.96)

0.78

(0.67-0.90)

0.56

(0.49-0.63)

3

0.58

(0.52-0.65)

0.55

(0.47-0.63)

0.37

(0.31-0.57)

4

0.31

(0.29-0.35)

0.30

(0.28-0.34)

0.25

(0.21-0.31)

5

0.30

(0.25-0.34)

0.29

(0.26-0.31)

0.27

(0.25-0.30)

Head (Fig. 8): Parietal sclerite with 6-7 setae in dorsolateral portion of sclerite.
Distal four palpal segment proportions: 1.0, 1.4, 1.4, 5.6. Antennal segment pro-
portions: 0.70, 0.72, 1.0, 0.76, 0.97, 1.01, 0.94, 0.96, 0.96, 0.99, 0.96, 0.98, 0.96,
0.88, 1.11; ultimate segment 1.3 length of penultimate.

Legs: Tibial spur formula 0-0-1. Progressive proportions of leg segment lengths
as below:

foreleg: 1.00, 0.96, 0.54, 0.42, 0.70, 0.55, 0.95
midleg: 1.00, 0.81, 0.52, 0.49, 0.70, 0.55, 0.94
hindleg: 1.00, 0.93, 0.38, 0.33, 0.66, 0.68, 1.06

Genitalia (Fig. 28): IXth tergite lobes very prominent, slightly convergent; lobe
shape irregular, outer comer acutely rounded, posterior margin oblique, projecting

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posteriorly as an angulate lobe, inner margin strongly inflated mesally, i.e. strongly
curved to a narrowed base; bristles numerous and regular over entire dorsal surface.
Tegmen entire. Paramere broad basally, tapering to distal four-fifths where abruptly
narrowed (outer wall terminating); apex acute and incurved; aperture subapical, at
termination of outer wall. Penis filaments 3 in number, equal in length, shorter than
parameres, all simple, truncate rods, median much heavier basally than laterals.

REMARKS

This is the largest species of the Tenuipes Group, although of average size for
the genus. In addition to diagnostic characteristics found in the genitalia of both sexes
(see keys), the adults appear distinctive, even to the naked eye, because of the light
brown prescutellar quadrate area of the scutum. The scuta of all the other species are
more or less unicolorous.

DISTRIBUTION (Fig. 42)

B. williamsae is known only from the extreme southern portion of the Appala-
chians at the confluence of Georgia, Tennessee and the Carolinas.

MATERIAL

Adults. — GEORGIA, Lumpkin County. De Soto Falls State Park, upper falls, 28 April
1973, H. D. Pratt (5 6 :HDP). NORTH CAROLINA, Macon County. Highlands, 3-5000 feet,
April-May 1936, R. C. Shannon (18 6 , 17 9, 4 pupae:USNM). Clear Creek, Highlands, 3200
feet, 1-2 July 1958, J. G. Franclemont (19 9:CU). County unknown. Bubbling Spring Creek,
5100 feet, 17 July 1957, J. G. Chillcott (1 $ :CNC). TENNESSEE, Sevier County. Great Smoky
Mountains National Park, Greenbrier Cove, 2000 feet, 18 May 1957, W. R. M. Mason (2
9:CNC); same locality, 2-2500 feet, 22 April 1939, 15 May 1938, I. W. Williams (<3 AL-
LOTYPE, 2 6 , 3 9 PARATYPES:USNM); upper Greenbrier Cove, slopes Mount Leconte,
4200 feet, 5 June 1939, C. P. Alexander (9 HOLOTYPE:CPA).

Pupae (pharate adults dissected). — UNITED STATES. TENNESSEE, Sevier County.
Greenbrier Cove, Great Smoky Mountains National Park, 22 May 1938, A. C. Cole (14 pu-
pae:UT). Oconaluftee River near Alum Cave Bluffs, Great Smoky Mountains National Park, 8
July 1974, D. & M. Davis (10 pupae:USNM).

TAXONOMY -PUPAE

The difficulties of distinguishing pupae because of their similarity among all the
species (excepting species F) was mentioned earlier. Size may be used when there is
reason to believe the sample contains disparate forms. Otherwise, since length-width
ratios are constant and length is a continuum from the smallest males to the largest
females, one must dissect the pharate adults to identify pupae. (Size data on samples
of known species are given in Table I).

DESCRIPTION

General. — Typical in basic structure for genus Blepharicera. Outline shape
ovate, cross section hemi-oval. Size. — 6 3. 6-4. 7; 9 4.7-5. 7 mm. Color (in life).
— Upper surface evenly black; lower surface white. Structure. — Dorsal cuticle of
metascutal, scutellar and abdominal sclerites, as well as small area immediately an-

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Table I

Blepharicera, Tenuipes Group. Size of pupae
(mean lengths in mm). N.a. = not available.

d

.9

B. capitata

n.a.

n.a.

B. cherokea

n.a.

n.a.

B. diminutiva

n.a.

n. a.

B. similans

3.62

4.70

B. tenuipes

4.37

5.04

B. williamsae

4.74

5.65

B. species F

n.a.

4.7

terior to branchial lamellae on branchial sclerite, finely and densely papillate; re-
mainder of dorsum smooth or weakly wrinkled. Branchiae parallel, widely separated;
posture erect, composed of 4 elongate, apically rounded, rigid plates; all plates of
equal length; internal pair more weakly sclerotized than external. Antennal cases of
both sexes extending to less than half the length of wing cases.

The material for this project includes one specimen which deviates from all the
others in the form of the tergites of abdominal segments I-V. These sclerites, instead
of forming a flat arch across the midline, possess a low, cone-shaped prominence
anteromedially; a similar, but lower and broader prominence arises from the scutellar
sclerite. The processes are presumedly homologous to those on the mid dorsum of
“Larva F.”

TAXONOMY-LARVAE

Six types of larvae are distinguishable in the material assembled for this study.
Because none could be definitely associated with adults, their identities are tentative
and in 3 cases still so uncertain that they can only be guessed.

DESCRIPTION

General. — Typical in basic structure for genus Blepharicera . Size. — Body
length (prepupal) 5-7 mm. Width of head capsule 1.2-1. 5 mm. Color. — Pigmentation
usually pale, yellowish, sometimes medium brown. Sclerotized structures medium to
dark brown. Structure. — Intercalary segments present. Head capsule with complete
incision in lateralia. Antennae very short and with two sclerotized rings. Dorsal pseu-
dopods absent. No dorsal spines, sclerotized plates or processes of any kind present
(one species with middorsal coniform swellings). Ventrolateral gills with 7 filaments
(6 occasionally on terminal division). Chaetotaxy . — Integument dorsally set with nu-
merous secondary, coniform or claviform sensilla, primary sensilla recognizable as
follows: tp M-T, members of pairs disjunct; ss P-T, that of P out of line laterad with
those of M-T; st, inner tp, and dpod I- VII.

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Larva A

(Tentatively B. tenuipes )

DESCRIPTION

Fourth instar larva (Fig. 30): Body length 6.9 mm; head width 1.48 mm. Sec-
ondary dorsal sensilla short, elongate-claviform; arranged in two irregular transverse
rows on trunk divisions and abdomen I; a single row on each thoracic segment; rows
confluent and sensilla general laterally, some setiform sensilla mixed. Recognizable
primary sensilla all setiform except t P-M which are subchaetiform.

MATERIAL (Fig. 41)

Larvae (and accompanying immature pupae) . — CANADA . NEWFOUNDLAND. Baie
Verte River, 23 June 1971, S. W. Frost (4 larvae:CNC). ONTARIO, Carleton County. Cun-
ningham and Riopelle Islands, Remic Rapids, 18 May, 6 June 1949 (3 pupae, 14 larvae:CNC).
Renfrew County. Petawawa, rapids belong Highway 17 bridge, 28 April 1959, G. E. Shewell
(5 pupae, 3 larvae:CNC). Timiskaming County. Kelly Creek, Highway 101, 28.8 mi W Timmins,
23 May 1972, ROM field party No. 720183 (14 larvae:ROM). Wentworth County. Spencer
Creek, below falls, Ontario Railway Station, Dundas, 13 May 1954 (2 pupae, 6 larvae:USNM).
QUEBEC, Saguenay County. Baie-Comeau, 11 June 1963, H. C. Connell (15 larvae:CNC).
STATE and County UNKNOWN. Bug River Rapids, 21 June 1951 (8 pupae, 5 larvae:USNM).
UNITED STATES. MARYLAND, Frederick County. Catoctin Creek, 1 mi N Little Catoctin,
Harmony, 30 May 1958, P. H. Freytag (1 larva:OSU). Garrett County. Savage River at Big
Run Campground, 17 April 1968, Yamamoto & Odum (1 larva:ROM).

REMARKS

Determination of this larva seems fairly certain because of its wide occurrence
in areas occupied exclusively by tenuipes. A significant locality in this respect is
Ithaca, New York, where the numerous collections of all stages have never contained
any deviant forms.

Larva B

(Tentatively B. similans)

DESCRIPTION

Fourth instar larva (Fig. 33): Body length 5.1 mm; head width 1.22 mm. Sec-
ondary dorsal sensilla short, extremely capitate (almost spherical in some), the largest
apically dentate; arranged in two irregular transverse rows on trunk divisions and ab-
domen I; a single row on each thoracic segment; rows confluent laterally and where
sensilla setiform mixed and general. Usually one conspicuously large, secondary clav-
iform sensillum in dorsolateral area. Recognizable primary sensilla all setiform.

MATERIAL (Fig. 40)

Larvae (and accompanying immature pupae). — CANADA. QUEBEC, Brome County. Sut-
ton Mountain Creek, 11 July 1929, G. S. Walley (19 larvae:CNC). UNITED STATES. MAINE,
Somerset County. Pleasant Pond Stream, Caratunk, 24 June 1966, R. & D. Koss (78 larvae:JH).
MARYLAND, Garrett County. “CCXXXIV falls, muddy creek”, 25 August 1965, J. Glime
(1 larva:USNM). MASSACHUSETTS, Berkshire County. Bashbish Creek, 6 July 1950, A.
Stone (4 pupae, 12 larvae:USNM). NEW HAMPSHIRE, Coos County. Creek on way to Glen

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House, 18 July 1929, G. S. Walley (1 pupa, 3 larvae:CNC). NORTH CAROLINA, Jackson
County. 3 mi E Balsam, 3400 feet, 11 July 1974, D. & M. Davis (12 larvae:USNM). Swain
County. Deep Creek at Deep Creek Campground, Great Smoky Mountains National Park, Bryson
City, 21 May 1970, Wiggins & Yamamoto No. 700365 (10 larvae:ROM). TENNESSEE, Sevier
County. Gatlinburg, fork Little Pigeon River, 27 May 1934, T. H. Frison (3 larvae:INHS).
Greenbrier Cove, Great Smoky Mountains National Park, 18 May 1952, H. B. Reed (53 lar-
vae:UT). VIRGINIA, Giles County. Big Stony Creek, Kimballton, 26 May 1962, C. V. Coveil
(1 pupa, 3 larvae:UL). Sinking Creek, 29 May 1941, A. Stone No. Ill (6 pupae, 10
larvae:USNM). Highland County. Cowpasture River, 3 mi N Williamsville, 27 July 1974, M.
M. Davis (1 pupa, 11 larvae:USNM). Madison County. White Oak Creek, Shenandoah National
Park, 24 June 1951, B. O. Burks (8 larvae:USNM).

REMARKS

Identification of this form is fairly certain and based upon several widely spaced
collections of larvae accompanying pupae with clearly recognizable pharate adults
(see MATERIAL under account of similans ).

Larva C

(Species Undetermined)

DESCRIPTION

Fourth instar larva (Fig. 35): Body length 6.4 mm; head width 1.33 mm. Sec-
ondary dorsal sensilla long, attenuo-claviform; arranged generally over each division
but tending to form transverse rows especially on thoracic segments; also fewer across
trunk segments slightly posterior to a mid-transverse line; laterally mixed with seti-
form sensilla. Recognizable primary sensilla all setiform.

MATERIAL (Fig.43)

Larvae (and accompanying immature pupae). — UNITED STATES. GEORGIA, Lumpkin
County. Hightower River, 11 April 1941, A. Stone No. 21 (5 larvae:USNM). Union County.
Vogel State Park, 11 April 1941, A. Stone No. 18 (9 larvae:USNM). MARYLAND, Frederick
County. Cunningham Falls State Park, 15 April 1973, P. & P. Spangler (23 larvae:USNM).
Garrett County. Ginsend Run, 4 May 1965, J. Glime (2 larvae:MSU). Sang Run Crossing, Sang
Run Road, 17 April 1968, Yamamoto & Odum (6 larvae:ROM). VIRGINIA, Alleghany County.
Tributary Jackson River beside Route 220 just N Covington, 18 April 1968, Yamamoto & Odum
(1 larva:ROM). Potts Creek, 3 mi SW Jordan Mines, 18 April 1968, Yamamoto & Odum (9
larvae:ROM). Amherst County. Tributary of James River beside Blue Ridge Parkway near junc-
tion with Route 501, 19 April 1968, Yamamoto & Odum (3 larvae:ROM). Bath County. Stream
in Blowing Springs Recreation Area on Route 39, 18 April 1968, Yamamoto & Odum (6 lar-
vae:ROM). Floyd County. Little Indian Creek, Copper Valley, 24 April 1966, P. & B. Wy-
godzinsky (6 larvae:AMNH). Giles County. Big Stony Creek, Kimballton, Station 2, 14 April,
5 May 1962, C. V. Covell (4 pupae, 25 larvae:UL). Sinking Creek, 29 May 1941, A. Stone
No. Ill (1 larva:USNM). Green County. Swift Run, 5 mi W Standardsville, Highway Route
33, 17 May 1970, R. W. Koss (10 pupae, 15 larvae:JH). Lydia, 20 April 1938, R. Burks (3
larvae:INHS). Madison County. Oak Creek, Shenandoah National Park, 6 May 1951, B. D.
Burks (5 larvae:USNM). White Oak Canyon, Skyline Drive, 6 May 1951, Burks & Sommerman
(2 larvae:USNM). Rappahannock County. Thornton River, Shenandoah National Park, 15, 20,
26 April 1961, O. & R. Flint (1 pupa, 29 larvae:USNM). Shenandoah County. Passage Creek,
Camp Roosevelt, 21 May 1973, S. Fiance (3 larvae:USNM). STATE and County UNKNOWN.
Treasler Run, Aitkin Farms, 29 April 1951, S. W. Frost (9 larvae:PSU).

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REMARKS

The distribution of this form coincides approximately with that of the atypical
type of tenuipes (see VARIATION under that species) and may represent its larva.
If this proves to be the case, its distinctiveness and constancy detract from the hy-
pothesis of hybrid origin for that type and suggest that a separate species or subspecies
is present.

Larva D

(Tentatively B. williamsae )

DESCRIPTION

Fourth instar larva (Fig. 31). — Body length 6.7 mm; head width 1.43 mm. Sec-
ondary dorsal sensilla short, lanceolate; arranged generally over each division but
tending to form transverse rows on thoracic segments; mixed with chaetiform sensilla
only near lateral margin. Recognizable primary sensilla all setiform except t P-I and
st V-VI which are cylindriform. Specimens are often more darkly pigmented dorso-
mesally than laterally and present a longitudinally banded appearance.

MATERIAL (Fig. 42)

Larvae (and accompanying immature pupae). — UNITED STATES. GEORGIA, Dawson
County. Amicalola Falls, 11 April 1941, A. Stone (2 larvae:USNM). Gilmer County. Little
Turnip Town Creek, Walnut Mountain Development, 13 March 1974, H. D. Pratt (6 pupae, 6
larvae :HDP). Rabun County. Tallulah River at Tallulah River Campground, S Tate City, 17
May 1970, Wiggins & Yamamoto (1 larva:ROM). Union County. Chestatee River, 11 April
1941, A. Stone (1 larva:USNM). NORTH CAROLINA, McDowell County. Woodlawn, 23
March 1940, Frison, Mohr & Hawkins (3 larvae:INHS). Swain County. New Found Gap, 3560
feet, 28 May 1934, T. H. Frison (6 pupae, 6 larvae:INHS). SOUTH CAROLINA, Oconee
County. Howard Creek, 6 February 1976, R. L. Green & D. A. Braatz (5 larvae:LACM). TEN-
NESSEE, Sevier County. Campground near Chimneys, Great Smoky Mountains National Park,
11 May 1944, Frison & Ross (16 larvae:INHS). Stream 10.2 mi E Gatlinburg entrance on Route
441, 20 May 1970, Wiggins & Yamamoto No. 700361 (10 larvae:ROM). Greenbrier Cove,
Great Smoky Mountains National Park, 13 February 1938, A. C. Cole (47 larvae:UT); 18 March
1939, A. C. Cole (2 larvae:INHS). Small stream 0.8 mi W New Found Gap on Route 441, 20
May 1970, Wiggins & Yamamoto No. 700359 (2 larvae:ROM).

REMARKS

Coincidence of large size and larval collections with the adults of williamsae
strongly suggest the identity of that species with this form.

Larva E

(Species Undetermined)

DESCRIPTION

Fourth instar larva (Fig. 32). — Body length 5.9 mm; head width 1.64 mm. Sec-
ondary dorsal sensilla all setiform (except for occasional claviform shapes on thoracic
segments in some specimens); arranged in two irregular rows on trunk divisions and
abdomen I; a single row on each thoracic segment; rows confluent and sensilla general

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laterally. Recognizable primary sensilla all setiform and similar to secondary sensilla
except for larger size.

MATERIAL (Fig. 44)

Larvae. — CANADA. ONTARIO, Parry Sound County. Cashman Creek crossing Highway
518 at Sand Lake, 1 June 1972, ROM field party, No. 720212 (1 larva:ROM). UNITED STATES .
MINNESOTA, Lake County. Split Rock Creek, 2 mi W Lake Superior, 16 May 1952, J. Ger-
berich (6 larvae:OSU). VIRGINIA, Giles County. Big Stony Creek, Station 2, Kimballton, 5
May 1962, C. V. Coveil (1 larva:UL). STATE and County UNKNOWN. Stillwater Creek, 13
June 1961 (1 larva:USNM).

REMARKS

I have no idea of the identity of this form. It is sympatric only with tenuipes,
but since the larva of that species is known, larva E would appear to belong to an
unknown entity.

Larva F

(Species Undetermined)

DESCRIPTION

Fourth instar larva (Figs. 34, 36). — Body length 5.8 mm; head width 1.25 mm.
A conspicuous median conical protuberance on each trunk division and abdomen I.
Posterolateral lobes of terminal division poorly developed, terminal incision very shal-
low. Secondary dorsal sensilla claviform; arranged generally on all divisions except
for 4-6 larger sensilla grouped tightly at apex of each coniform process; lateral sensilla
mostly setiform. Recognizable primary sensilla well marked, all setiform except for
t P-T and t V-VI which are cylindriform.

MATERIAL (Fig. 44>

Larvae (and accompanying immature pupa). — CANADA. NEW BRUNSWICK, Charlotte
County. Saint Stephen, 27 June 1965, G. E. Shewed (1 pupa, 2 larvae:CNC). UNITED STATES.
GEORGIA, Lumpkin County. Hightower River, 11 April 1941, A. Stone No. 21 (11
larvae:USNM). MAINE, Somerset County. Pleasant Pond Stream, Caratunk, 24 June 1966, R.
& D. Koss (1 larva:JH). NORTH CAROLINA, Avery County. Linville River at Linville Falls
Campground, Blue Ridge Parkway, 22 May 1970, Wiggins & Yamamoto No. 700369 (11 lar-
vae:ROM). Swain County. Deep Creek at Deep Creek Campground, Great Smoky Mountains
National Park, Bryson City, 21 May 1970, Wiggins & Yamamoto No. 700365 (2 larvae:ROM).
Noland Creek, Great Smoky Mountains National Park, 24 May 1974, R. L. Green (1 larva:LACM).
VIRGINIA, Bath County. Stream in Blowing Springs Recreation Area on Route 39, 18 April
1968, Yamamoto & Odum (1 larva:ROM).

REMARKS

Unfortunately the female dissected from the single pupa available of this form
was immature and not identifiable. Because of its great latitudinal range it seems
unlikely that it belongs to either capitata, cherokea or diminutiva, all species now
known to have restricted distribution. Possibly another new species is indicated.

1

CHARACTERS for explanation of abbreviations.

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Figures 3-8: Anterior view of head capsules of male Blepharicera: 3. B. capitata. 4.
B. cherokea. 5. B. diminutiva. 6. B. similans. 7 . B. tenuipes. 8. B. williamsae.

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Blephariceridae Eastern North America

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10. B. cherokea. 11. B. diminutiva. 12. B. similans. 13. B. williamsae.

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Figures 14-15: Terminology of head capsule, Blepharicera tenuipes , female: 14. An-
terior view. 15. Posterior view, mouthparts omitted.

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Blephariceridae Eastern North America

35

Figures 16-21: Male (left) and female (right) genitalia of Blepharicera: 16-17. B.
capitata. 18-19. B. cherokea. 20-21. B. diminutiva.

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Figures 22-29: Male (left) and female (right) genitalia of Blepharicera (continued):
22-23. B. similans. 24-27. B. tenuipes (24. Ninth tergite lobes of “southern type”;
25. Three views of apex of paramere of male genitalia). 28-29. B. williamsae.

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Blephariceridae Eastern North America

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Figures 30-36: Fourth instar larvae of Blephar ic era, 34. lateral view of Larva F (spe-
cies undetermined), remainder dorsal view of left side of third abdominal segment
showing details of chaetotaxy. 30. Larva A (probably tenuipes ). 31. B. williamsae.
32. Larva E (species undetermined). 33. Larva B (probably similans ). 35. Larva C
(species undetermined). 36. Larva F (species undetermined).

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Figures 37-40. Distribution of Blepharicera in eastern North America. Points indicate
Joci of collections, line encloses probable total range. 37. Typical B. capitata (dots)
and “separata” type (squares). 38. B. diminutiva. 39. B. cherokea. 40. B. similans
adults (dots) and larvae (triangles).

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Blephariceridae Eastern North America

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Figures 41-44. Distribution of Blepharicera in eastern North America (continued).
Points indicate loci of collections, line encloses probable total range. 41. Typical
B. tenuipes adults (dots), southern type (squares) and larvae (triangles). 42. B. wil-
liamsae adults (dots) and larvae (triangles). 43. Larva C. 44. Larva E (triangles and
larva F (squares).

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No. 291

ACKNOWLEDGMENTS

This study is based on 1869 specimens (773 adults, 1096 larvae and pupae). For
the use of this material I wish to acknowledge the following repositories (listed al-
phabetically by the acronym used in citing specimens in the text above) and thank
their respective curators who kindly arranged loans and provided much information:
AMNH - American Museum of Natural History, Pedro Wygodzinsky.

ANSP - Academy of Natural Sciences of Philadelphia, Harold Grant (deceased).

BMNH - British Museum, Natural History, Paul Freeman.

CAS - California Academy of Sciences, Paul H. Arnaud.

CNC - Canadian National Collection, B. V. Peterson.

CPA - Private collection of Charles P. Alexander, Amherst, Massachusetts.

CU - Cornell University, L. L. Pechuman.

DPC - Duke Power Company, Environmental Laboratories. R. Lynn Green.
HDP - Private collection of Harry D. Pratt, Atlanta, Georgia.

INHS - Illinois Natural History Survey, R. T. Allen.

JH - Johns Hopkins University, Richard Koss.

KS - Kansas State University, Norman Marston.

KU - University of Kansas, Snow Entomological Museum, George W. Byers.
LACM - Natural History Museum of Los Angeles County.

MCZ - Harvard University, Museum of Comparative Zoology, Philip J.

Darlington.

MMU - McMaster University.

MSU - Michigan State University, Roland L. Fischer and Frederick W. Stehr.
OS - Oregon State University, J. D. Lattin.

OSU - Ohio State University, Paul Freytag.

PSU - Pennsylvania State University.

ROM - Royal Ontario Museum, Glenn B. Wiggins.

UCR - University of California, Riverside, Saul Frommer.

UL - University of Louisville, Charles V. Covell, Jr.

USNM - U.S. National Museum of Natural History Collection, Alan Stone, F.

Christian Thompson, and W. W. Wirth, Agricultural Research Service,
United States Department of Agriculture.

UNH - University of New Hampshire, J. G. Conklin.

UT - University of Tennessee, Arthur C. Cole, Jr.

I wish also to express my great appreciation for the help generously donated by
both Charlotte Low (preparation of specimens and compilation of statistical data) and
Carol Madle (assistance with illustrations and rendering of figures 12-13). D. A. Craig
kindly made available specimens of first instar Neocurupira chiltoni.

Finally, I extend my thanks to my colleagues Julian P. Donahue, Roy R. Snelling
and members of the review and editorial committees for criticism of the manuscript.

LITERATURE CITED

Aldrich, J. M. 1905. A catalogue of North American Diptera (or two-winged flies). Smith.
Misc. Colls. 46(1 444): 1-680.

1978

Blephariceridae Eastern North America

41

Alexander, C. P. 1953. Undescribed species of nematocerous Diptera. Part II. Brooklyn En-
tomol. Soc., B. 48:41-49.

1963. Blepharoceridae and Deuterophlebiidae in Guide to the insects of Connecticut.

Part VI. The Diptera or true flies. Fasc. 8, pp. 39-83.

Comstock, J. H. and A. B. Comstock. 1895. A Manual for the Study of Insects. Comstock
Pub. Co., Ithaca.

Craig, D. A. 1967. The eggs and embryology of some New Zealand Blephariceridae (Diptera,
Nematocera) with reference to the embryology of other nematocera. Roy. Soc. New Zea-
land, Trans. 8:191-206.

Hogue, C. L. 1973. The net-winged midges or Blephariceridae of California. Calif. Insect Sur-
vey, B. 15:1-83.

Hora, S. L. 1930. Biology, bionomics and evolution of the torrential fauna, with special ref-
erence to the organs of attachment. Roy. Soc. London, Phil. Trans. Ser. B. 218:217-282.

Johannsen, O. A. 1903. Aquatic nematocerous Diptera. New York State Mus., B. 68:328-
441.

1929. A new species of Blepharocera from Massachusetts (Diptera). Psyche 36:123-

124.

1934. Aquatic Diptera. Part I. Nemocera, exclusive of Chironomidae and Cerato-

pogonidae. Cornell Univ. Agric. Expt. Sta., Mem. 164:1-71, pis. I-XXIV.

Kellogg, V. L. 1900. Notes on the life-history and structure of Blepharocera capitata Loew.
Entomol. News 11:305-318.

1903. The net-winged midges (Blepharoceridae) of North America. Calif. Acad. Sci.,

Proc. 3rd series 3:187-233.

1907. Diptera. Fam. Blepharoceridae. Genera Insectorum 56:1-15.

Loew, H. 1863. Diptera americae septentrionalis indigena. Centuria quarta. Berliner Entomo-
logischer Zeitschrift 7:275-326.

Osten Sacken, C. R. 1895. Contributions to the study of the Liponeuridae Loew (Blepharo-
ceridae Loew, olim). Berliner Entomologischer Zeitschrift 40:148-169.

Riley, C. V. 1881. Notes on Blepharoceridae. Amer. Natur. 15:438-447, 748.

Stone, A. 1965. Family Blephariceridae. Pages 99-100 in Stone, A., C. W. Sabrosky, W. W.
Wirth, R. H. Foote and J. R. Coolson. A catalog of the Diptera of America North of
Mexico. U. S. Dep. Agr., Washington, D.C.

Vaillant, F. 1959. The larvae of three nearctic Diptera of the family Psychodidae. New York
Entomol. Soc., J. 67:39-50.

Walker, F. 1848. List of the specimens of dipterous insects in the collection of the British
Museum. 1:1-229.

Accepted for publication June 15, 1977.

NUMBER 292
MARCH 25, 1978

scn> 7$

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NEW SPECIES AND RECORDS OF TROPICAL WEST AMERICAN
MARGINELLIDAE (MOLLUSCA: NEOGASTROPODA)

By Barry Roth

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

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NEW SPECIES AND RECORDS OF TROPICAL WEST AMERICAN
MARGINELLIDAE
(MOLLUSCA: NEOGASTROPODA)1

By Barry Roth2

Abstract: Six new species of Marginellidae are described: Prunum ( Prunum )
gorgonense from off Gorgona Island, Colombia; Prunum ( Prunum ) macleani from
Ecuador; Prunum ( Microspira ) aletes and Dentimargo zetetes from localities on the
Pacific coast of Costa Rica; Dentimargo epacrodonta from the Republic of Panama;
and Volvarina innexa from the Galapagos Islands. Prunum {Microspira) aletes is
similar to several Caribbean species and Neogene species from Florida. Prunum
{Prunum) curtum (Sowerby) ranges into temperate waters south of the Panamic prov-
ince; records from the Gulf of Panama need confirmation. The radula of Prunum
{Prunum) woodbridgei (Hertlein and Strong) is illustrated and compared to that of
Prunum storerium (Couthouy). Dentimargo anticlea (Dali) is tentatively recognized
from several Galapagan localities.

INTRODUCTION

Recent accounts of the members of the Marginellidae known from the Panamic
faunal province of tropical western America (Coan and Roth in Keen 1971; Roth and
Coan 1971) cite 20 named species and subspecies. The extensive molluscan collec-
tions of the Section of Malacology of the Natural History Museum of Los Angeles
County (LACM), and the gastropod collections of the Allan Hancock Foundation,
currently on loan to that section, have furnished material for the description of the
following new species and additional information concerning species previously
described.

I acknowledge the courtesy of James H. McLean, Curator of Invertebrate Zo-
ology, LACM, in making these collections available for study and aiding in the prep-
aration of this paper. For assistance of various kinds during this investigation I am
indebted to Emily H. Vokes, Eugene V. Coan, Antonio J. Ferreira, and Patrick
LaFollette. An earlier draft of part of the manuscript was read critically by J. Wyatt
Durham. Joseph Rosewater, Thomas R. Waller, and Frederick J. Collier lent com-
parative material from the National Museum of Natural History. Phillip W. Clover
supplied information on type material studied by him in European museums.

Review Committee for this Contribution
William K. Emerson
James H. McLean
Joseph Rosewater

department of Geology, California Academy of Sciences, San Francisco, California 94118.

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FAMILY MARGINELLIDAE FLEMING

GENUS PRUNUM HERRMANNSEN 1852

Type species, by monotypy and tautonymy, Voluta prunum Gmelin 1791; Recent, Caribbean

region.

As stated by Woodring (1970), Prunum is the proper generic allocation for a
large number of fossil and Recent species. Prunum in the broad sense includes ovate
to elongate shells with medium to low spire, aperture narrow or wide but not flaring
anteriorly, outer lip more or less thickened and smooth or denticulate, and some de-
velopment of callus around the aperture.

Subgenus Prunum sensu stricto

Outer lip not greatly thickened, smooth within; anterior canal shallow; callousing
around aperture generally limited to small patch on parietal wall; color pattern, when
present, generally of indistinct banding.

Prunum ( Prunum ) gorgonense new species
Figure 1

Diagnosis ; A small, elongate Prunum differing from other west American species
in the relatively high spire, narrow body whorl, and short aperture, colored pale pink-
ish tan with interior of aperture yellow.

Description of holotype: Shell rather small for the genus, elongate-ovate, nar-
rower anteriorly, highly polished, solid. Color pale pinkish tan with two very faint,
narrow, darker spiral bands dividing the body whorl into thirds, and a narrow zone
of apricot-orange immediately below the suture. Apex translucent, apricot-orange;
outer lip white; interior of aperture golden yellow. Spire elevated, apical angle about
60°, covered by a transparent glaze, which leaves sutures visible but impalpable. Outer
lip with simple convex curvature, sharply varicose externally, not denticulate. Ap-
erture narrow, wider anteriorly, with a short posterior notch; anterior margin evenly
rounded, without indentation. Parietal wall moderately convex, not calloused. Col-
umella very slightly concave, with four oblique folds including fold at base of col-
umella, the two most anterior folds close together and fusing distally into a low, white
fasciole that proceeds around anterior end of shell to merge with outer lip callus.
Length 16.2 mm, width 8.0 mm.

Type locality: AFH 221-34, off Gorgona Island, Colombia (3° OF 25" N, 78°
10' W), in 37 m on rock and shell bottom. Collected by Allan Hancock Pacific Ex-
pedition of 1934, 12 February 1934. One specimen.

Type material. Holotype, LACM-AHF 1777.

Referred material: One immature specimen, AHF 228-34, between Gorgona and
Gorgonilla Islands, Colombia (2° 56' 20" N, 78° 14' W), mud and sand bottom. Col-
lected by Allan Hancock Pacific Expedition of 1934, 12 February 1934.

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West American Marginellidae

3

Discussion: Prunum ( Prunum ) gorgonense is smaller and much paler than
Prunum (. Prunum ) sapotilla (Hinds 1844), which is grayish brown with a deep brown
aperture, tinged with dull orange outside the outer lip. The new species has a pro-
portionally higher spire, shorter aperture, and less inflated body whorl than P. sa-
potilla. The latter species appears to be confined to the region of the Gulf of Panama.

In shape P. gorgonense is closer to Prunum {Prunum) godini Olsson 1964, from
the Esmeraldas Formation, Neogene of Ecuador, and may be a descendant of that
species (Figs. 2, 3). Prunum godini is larger (holotype 21.8 mm long), broader at the
shoulder, and has the columella markedly excavated anteriorly so that the two most
anterior folds diverge, rather than converge as in P. gorgonense . The fossil species
also has a raised vertical callus rim anteriorly, visible in left side view (Fig. 3), which
is absent in P. gorgonense.

The columellar folds of P. gorgonense like those of other species of the subgenus
are concave in profile, with the hollow surface directed posteriorly.

Etymology: gorgonense — from Gorgona Island, the type locality.

Prunum {Prunum) curtum (Sowerby 1832)

Figure 4

Marginella curta Sowerby 1832:105; Reeve 1864: pi. 6, figs. 23a, 23b.

Marginella {Prunum) curta Sowerby, Coan and Roth 1966:280, pi. 48, figs. 4-6.

Prunum {Prunum) curtum (Sowerby), Coan and Roth in Keen 1971:633, fig. 1398.

Although it was described from Iquique, Chile, and Paita, Peru, few collections

Figures 1-3. 1. Prunum {Prunum) gorgonense new species, holotype, FACM-AHF
1777, length 16.2 mm. 2-3. Prunum {Prunum) godini Olsson, holotype, USNM
643945, length 21.8 mm.

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Contributions in Science

No. 292

of this species have been made south of the Panamic province. Fresh beach shells
were found in some numbers on an intertidal sandbar at Laguna Grande, lea Province,
Peru (14° 08' S, 76° 15' W) by J. H. McLean and Victor Alamo, 31 March 1972.
One specimen from that locality (LACM 72-77) is illustrated. An analogous situation
in the Atlantic Ocean is the presence of Prunum species in temperate waters of Ar-
gentina (Carcelles 1944).

No verified type rpaterial of this species is known to exist (Coan and Roth 1966).
The specimens from Laguna Grande agree with the original diagnosis and with
Reeve’s (1864) figure of a specimen from the Cuming collection. For purposes of
comparison with Prunum {Prunum) macleani, I have considered the Laguna Grande
material to represent typical P . curtum.

Coan and Roth in Keen (1971) cite this species as ranging north to Manta, Ec-
uador. Lots at LACM from the Gulf of Panama differ in several details from South
American specimens and may represent other, unrecognized, species.

Prunum ( Prunum ) macleani new species
Figures 5-8

Diagnosis: A large Prunum with relatively low spire, differing from other west
American species in the elongate-conic body whorl, wide aperture, and coloration:
muted rose with white flecking in spiral bands, apertural callus edged with bright
orange.

Description ofholotype: Shell large for the genus, inflated, ovate-conic, narrower
anteriorly, highly polished, solid. Color muted rose to rosy tan with a lighter, poorly
defined, flesh color subsutural band and fine white flecking tending toward arrange-
ment in spiral bands, most evident around middle of body whorl and near anterior
end. Apex translucent, rosy brown; spire low, apical angle about 80°, light flesh color,
covered by a transparent glaze which renders suture indistinct. Aperture moderately
wide, wider anteriorly, rose color within, anterior margin somewhat effuse. Outer lip
varicose, white on ventral surface, margined with brilliant orange externally, not den-
ticulate. Parietal wall moderately convex with a patch of translucent, whitish callus.
Columella concave, with four oblique, moderately diverging folds, including fold at
base of columella; the two most anterior folds most oblique, nearly parallel, and closer
together than posterior two folds, merging distally into a low, white fasciole with a
bright orange external margin that proceeds around anterior end of shell to merge with
outer lip callus. Length 36.6 mm, width 21.2 mm.

Type locality: Station 778, Anton Bruun Cruise 18B (LACM 66-198), west of
Cabo Pasado, Ecuador (0° 21' S, 80° 41' W), in 19 m, collected 12 September 1966.
Twelve specimens.

Type material: Holotype, USNM 749065. One paratype, USNM 749066. Nine
paratypes (shells) and one paratype radula slide, LACM 1778. One paratype, Cali-
fornia Academy of Sciences, Geology Type Collection 55607. The paratypes differ
slightly from the holotype in height of spire, posterior extent of outer lip, degree of
development of parietal callus, and depth of ground color. In the lowest- spired ex-

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West American Marginellidae

5

Figures 4-7. 4. Prunum ( Prunum ) curtum (Sowerby), hypotype, CASG 54324, La-
guna Grande, lea Province, Peru, length 17.8 mm. 5-6. Prunum ( Prunum ) macleani
new species, holotype, USNM 749065, length 36.6 mm. 7. Prunum {Prunum) ma-
cleani new species, hypotype, AHF 213-34, off La Plata Island. Ecuador, length 22.7
mm.

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No. 292

ample, the apical angle is about 130° and the outer lip extends nearly as far backward
as the tip of the spire.

Referred material: Thirteen specimens, AHF 213-34, off La Plata Island, Ec-
uador (1° 15' 25" S, 81° 05' 15" W), 13-18 m on rock bottom. Collected by Allan
Hancock Pacific Expedition of 1934, 10 February 1934. Three specimens, AHF 212-
34, off La Plata Island, Ecuador (1° 15' S, 81° 04' 15" W), 82-100 m on rock and
mud. Collected by Allan Hancock Pacific Expedition of 1934, 10 February 1934.
These specimens range from 15.4 to 23 mm in length and are more strongly calloused
on the face of the body whorl (Fig. 7).

Discussion: This handsome species is the largest marginellid known from the
eastern Pacific region and, in the form represented by the type lot, could not be con-
fused with any other species. The referred material from off La Plata Island, however,
is much smaller and superficially resembles Prunum ( Prunum ) curtum. These spec-
imens are distinguished from P. curtum by their roseate color, more elongate-conic
body whorl, and effuse anterior end of the aperture. Prunum curtum is usually grayish
yellow or grayish brown with a variable amount of lighter flecking. Its body whorl
is shorter and broader than that of P. macleani, with a characteristic gibbous angu-
lation of the upper part. Its anterior end appears truncated in ventral view. The exterior
of its outer lip callus is brownish orange, and this same color appears, often quite
extensively, around the edge of the parietal callus (Fig. 4). The interior of the aperture
is commonly brown.

The radula of Prunum {Prunum) macleani (Fig. 8) consists of broad, nearly
straight, comblike rachidian teeth on which large and small cusps alternate. Each tooth
of the figured specimen bears 29 cusps.

Etymology: The species is named for Dr. James H. McLean.

Prunum {Prunum) woodbridgei (Hertlein and Strong 1951)

Figure 9

Marginella woodbridgei Hertlein and Strong 1951:80, pi. 26, figs. 3, 4.

Prunum {Prunum) woodbridgei (Hertlein and Strong), Coan and Roth in Keen 1971:633, fig.

1400.

Figure 9 illustrates the radula of a specimen from AHF 930-39, off San Jose
Light, Guatemala (13° 52' 35" N, 91° 01' 02" W), 22-24 m on fine black sand, col-
lected 23 March 1939. The radula consists of a single row of approximately 35 broad,
straight, comblike rachidian teeth on which large cusps alternate with from 1-4 smaller
cusps. A total of 24 cusps are present on each tooth of the figured specimen.

The radula is closely similar to that of Prunum storerium (Couthouy 1837) (Figs.
10, 1 1), a Caribbean species. The shell characters noted by Hertlein and Strong (1951)
also suggest close relationship between the two species. The radula of Prunum api-
cinum (Menke 1828), suggested by Coan and Roth (1966) as another allied species,
is less similar. Prunum storerium frequently is placed in the subgenus Microspira
Conrad 1868, based on the extensive callousing around the aperture and on the face
of the body whorl {cf. Woodring 1970:331-332); but this character evidently cuts

1978

West American Marginellidae

7

Figures 8-9. 8. Prunum ( Prunum ) macleani new species, radula of paratype, LACM
1778. 9. Prunum {Prunum) woodbridgei (Hertlein and Strong), radula, AHF 930-39,
off San Jose Light, Guatemala.

Figures 10-11. Radulae of Prunum storerium (Couthouy), Margarita Island, Venezuela.

across phyletic lines based upon radular features, which are assumed to be more
conservative.

Subgenus Microspira Conrad 1868

Type species, by monotypy, Volutella (Microspira) oviformis Conrad 1868; Miocene, Virginia.

Outer lip thick, smooth to finely denticulate within; anterior canal shallow; spire low,
sometimes depressed but not fully involuted, covered with a wash of callus; callousing
generally extensive around aperture and on face of body whorl; color pattern, when
present, of banding and/or irregular spotting.

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No. 292

Pending further studies, Egouena Jousseaume 1875 (type species, Egouena
egouen Jousseaume = Marginella amygdala Kiener; Recent, west Africa), and J.ep-
tegouana Woodring 1929 (type species, Voluta guttata Dillwyn; Recent, West Indies),
are here tentatively considered synonyms of Microspira, an interpretation suggested
by Woodring (1970). The chief characters on which supraspecific taxa in the Prunum
group are traditionally recognized pertain to development of callus on body whorl and
spire and the degree of thickening of the outer lip — characters having limited phyletic
significance, as discussed above under Prunum C Prunum ) woodbridgei . With the ex-
clusion of the generally less calloused species of Prunum, (sensu stricto), Micros-
pira, as broadly construed, contains the majority of recent and fossil species of the
genus.

Prunum (Microspira) aletes new species
Figures 12-13

Marginella ( Prunum ) species, Coan and Roth 1966:280, pi. 48, figs. 7, 8.

Prunum ( Microspira ) species Coan and Roth, Woodring 1970:332.

Diagnosis: A pyriform marginellid distinguished from other west American spe-
cies of Prunum by its ground color of orange-brown with indistinct darker brown

Figures 12-13. Prunum ( Microspira ) aletes new species, holotype, LACM 1779,
length 15.2 mm.

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West American Marginellidae

9

spiral bands and an overall pattern of irregular white flecks, narrow, sinuous aperture,
denticulate outer lip, and calloused parietal wall.

Description ofholotype: Shell of medium size for the subgenus, solid, pyriform,
inflated at shoulder, narrower anteriorly, with a shallow constriction across body
whorl slightly anterior to middle. Entire shell somewhat dorsoventrally flattened. Sur-
face highly polished, unsculptured. Color pale orange-brown with three darker, rather
indistinct, purplish brown spiral bands, the first immediately below the suture, the
second at the middle of the body whorl, and the third at the anterior one-fourth of
body whorl. Shell completely overlain by a layer of clear enamel, rendering suture
indistinct and, on the body whorl, bearing numerous white flecks of irregular shape,
which tend to fall into ranks parallel to outer lip and into series oblique to axis of
shell. Spire low, light apricot-orange; extreme apex white. Outer lip white, sharply
varicose externally and bearing two brown spots aligned with the two lower color
bands on body whorl; slightly constricted medially; finely denticulate along most of
its length. Aperture narrow, even, shallowly S-curved, banded brown and apricot-
orange inside; terminating posteriorly in a deep, straight sulcus. Parietal wall concave
anteriorly, convex posteriorly, covered with translucent white callus that thickens to
rim the posterior sulcus and continues up apertural side of spire nearly to apex. Center
of parietal callus thickened into an acute, axially elongate node. Columella with four
nearly equal, diverging folds, including one at base of columella, the two lowest folds
terminating anteriorly in a weakly developed, white fasciole that is continuous with
the varicose outer lip around anterior end of shell. Length 15.2 mm, width 10.0 mm.

Type locality: LACM locality 72-21, 3-11 m in cove between Isla San Jose and
Isla Cocinero, Islas Murcielagos, Guanacaste Province, Costa Rica (10° 51' 50" N,
86° 55' 30" W), collected by P. LaFollette, D. Cadien, A. J. Ferreira, 17 February
1972 (R/V Searcher station 404-405). Four specimens.

Type material : Holotype, LACM 1779. Two paratypes, LACM 1780. One par-
atype, California Academy of Sciences, Geology Type Collection, 55608. The par-
atypes differ very slightly from the holotype in development of the white spotting on
the body whorl and in extent of the parietal callus. Two are of paler ground color
than the holotype, with light orange, rather than brown, transverse bands. The par-
atypes range from 13.7 to 16 mm in length.

Referred material: The species was taken by the R/V Searcher at eight other
stations off Costa Rica: 392, LACM 72-12, 1.6 km offshore between Bahia Elena
and Juanillo Bay, Guanacaste Province (10° 57' 20" N, 85° 46' 08" W), in 53.0-26
m, 14 February 1972, 1 immature specimen; 403, LACM 72-20, cove on west side
of Isla Cocinero, Islas Murcielagos, Guanacaste Province (10° 51' 27" N, 86° 55' 15"
W), in 12-18 m, 16 February 1972, one specimen; 409, LACM 72-24, anchorage in
cove, northeast side of Isla San Pedrita, Islas Murcielagos, Guanacaste Province (10°
51' 40" N, 86° 57' 54" W), in 18-24 m, 17 February 1972, two specimens; 415,
LACM 72-30, south tip of Punta Santa Elena, Guanacaste Province (10° 53' 35" N,
85° 57' 52" W), in 12-15 m, 18 February 1972, four specimens; 423, LACM 72-37,
between the two Vivadores Norte Islands, near Bahia Culebra, Guanacaste Province
(10° 36' 42" N, 85° 43' 00" W), in 18-21 m, 19 February 1972, two specimens; 428,

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LACM 72-40, 1.6 km off beach, Bahia Brasilito, Guanacaste Province (10° 25' 57"
N, 85° 49' 18" W), in 18 m, 20 February 1972, three specimens; 431-432, LACM
72-42, 2.4 km east of Punta Ballena, Bahia Ballena, Puntarenas (9° 44' 15" N, 84°
33' 45" W), in 3-15 m, 21 and 22 February 1972, one specimen; 480-481, LACM

72- 67, off Isla del Cano, NW side, Puntarenas Province (8° 45' N, 84° 00' W), in

73- 82 m, 16 March 1972, one specimen.

One specimen, AHF 245-34, off Pacora Island, off Bahia Honda, Panama (7°
44' 19" N, 81° 35' 23" W), 27-46 m on rock and shell bottom, collected 21 February
1934.

Two specimens, California Academy of Sciences, Geology Type Collection
12749, 12750, from CAS locality 27557, dredged off Puntarenas, Costa Rica, by the
Templeton Crocker Expedition, 1 July 1932. These two worn specimens are the ones
described by Coan and Roth (1966:280, pi. 48, figs. 7, 8) as “ Marginella {Prunum)
sp.”

Discussion: Placement of this species in the subgenus Microspira was first sug-
gested by Woodring (1970), citing the Coan and Roth (1966) illustrations. This is the
first verified record of the subgenus in west American waters. {Marginella albuminosa
Dali 1919, probably a Microspira, is now thought to have been reported from west
Mexico in error; cf. Keen 1971:907.)

The color pattern of Prunum {Microspira) aletes distinguishes it from all other
known eastern Pacific Marginellidae. Its pyriform shape, narrow, S-curved aperture,
and dorsoventral flattening of the shall are other diagnostic characters. Prunum sa-
potilla (Hinds), which may be sympatric withP. aletes in the region of Bahia Honda,
Panama, sometimes has faint spiral bands, but it is much more elongate and lacks any
trace of white spotting. Coan and Roth (1966: table 2) contrasted the new species to
Prunum curtum (Sowerby) on several features, including the narrow aperture, the
thick, finely toothed outer lip, and the outline of the shell. Prunum woodbridgei
(Hertlein and Strong) is brownish gray, not spotted, and has an open, simply curved
aperture.

The dark spiral bands of P . aletes are at least partially antecedent to the clear
enamel with white spots that lies over them. An immature specimen (LACM 72-12)
is plainly banded and has much sparser spotting than any of the adult specimens.
Young specimens of the Caribbean Prunum storerium (Couthouy) are conspicuously
banded with purple on a straw-colored ground, this pattern later being obscured by
the predominantly gray enamel of mature shells.

Like many other tropical west American marginellids, Prunum {Microspira) al-
etes is most similar to species now living in the Caribbean region. The actual number
of western Atlantic species remains to be determined. Probably closest to P. aletes
is the species from Grand Cayman Island, British West Indies, figured by Abbott
(1958:84, pi. 2, figs, j, k) as “ Prunum pruniosum Hinds” (error for Marginella prui-
nosa Hinds 1844) with Marginella nivea C. B. Adams 1850, placed in synonymy.
Abbott (1958:84) described the species thus: “Shell 8 to 12 mm. in length, resembling
guttatum, but with its white spots half as small; with numerous, weak, uneven, den-
ticulations on the inner [outer?] lip; with 3 weak, diffused spiral bands of yellowish
brown (or absent); without color spots on the outer lip, and with a slightly raised spire

1978

West American Marginellidae

11

which is never covered by the labral callus.” Some color variation among Grand
Cayman populations was also specified. The narrow aperture is doubly curved, as in
P. aletes.

Tryon (1882) united Marginella pruinosa and Mar ginella nivosa Hinds 1844,
under the latter name; this synonymy was disputed by Abbott (1958). Marginella
nivea C. B. Adams 1850, described from Jamaica, is also similar. The lectotype,
figured by Clench and Turner (1950), is a worn, whitish shell 9 mm long, of the same
general shape as Abbott’s Grand Cayman species, with columellar folds more nearly
parallel. Tryon (1882) cited M. nivea as another synonym of M. nivosa. Examination
of large suites of Caribbean material may be necessary to settle the question of syn-
onymy. Some workers consider the presence or absence of dark spots on the outer
lip to be taxonomically significant.

Prunum ( Microspira ) guttatum (Dillwyn 1817) from Florida, the West Indies,
and Panama, is another white-flecked species. It is larger than P . aletes, frequently
reaching 20 mm or more in length, and more regularly ovate than the Pacific species.
Its apertural callus may extend as far as, or even posterior to, the tip of its low spire.
The outer lip is slightly constricted medially and bears a variable number of orange-
brown blotches. The labral denticulation is very weak in all specimens examined. In
form and color, P. guttatum is quite variable; notes on one variation were given by
Bayer (1943).

The somewhat similar Prunum {Microspira) roscidum (Redfield 1860), living
from New Jersey to eastern Florida, was contrasted to P. guttatum by Abbott (1957).
In the synonymy ofP. roscidum, Abbott included Marginella eulima Dali 1892, late
Neogene of Shell Creek and Caloosahatchie beds, Florida, which was described as
having opaque white flakes on translucent enamel and was itself a renaming of spotted
specimens previously (Dali 1890) referred to Marginella limatula Conrad 1834. Dali
(1892) suggested that M. eulima might be ancestral to M. nivosa. Spotted patterns,
sometimes detectable under ultraviolet light, characterize a number of other small to
medium-sized species from the Neogene of Florida.

Prunum {Microspira) donovani (Olsson 1967) of the Pinecrest beds, middle Pli-
ocene of southern Florida (Akers 1974), more than twice the size of P. aletes and
having five instead of four columellar folds, appears to be another relative. Under
ultraviolet light it shows a pattern of spots in the superficial enamel and, beneath,
transverse banding (Olsson 1967: pi. 8, figs. 9a-9c).

Throughout the later Tertiary, Microspira constitutes a prolific group in the
American tropics. Discrimination of probable lineages is difficult. It seems certain,
however, that several stocks of white-spotted species existed in the area prior to the
closing of the Central American isthmus. On the west, one such stock has survived
to the present day, represented by P. aletes.

Etymology: aletes — Gr., a wanderer.

GENUS DENTIMARGO COSSMANN 1899

Type species, by original designation, Marginella dentifera Lamarck 1803; Eocene, western

Europe.

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Contributions in Science

No. 292

Small to minute, biconic, high-spired; anterior notch absent; outer lip smooth or
denticulate within, commonly with a large posterior denticle.

Eburnospira Olsson & Harbison 1953 (type species, Marginella eburneola Con-
rad), Longinella Laseron 1957, non Gros and Lestage 1927 (type species, Marginella
maugeana Hedley), and Volvarinella Habe 1951 (type species, V. makiyamai Habe),
are regarded as synonyms of Dentimargo .

Dentimargo anticlea (Dali 1919)

Marginella anticlea Dali 1919:307. Coan and Roth 1966:289, pi. 51, fig. 61 (in synonymy of

Volvarinella eremus [Dali]).

Coan and Roth (1966) synonymized this species with Marginella eremus Dali
1919, considering the type lot ofM. anticlea to represent immature specimens of M.
eremus. The lectotype of anticlea is smaller, less sharply biconic, and lacks the prom-
inent labral denticle of M. eremus. Its nucleus is moderately large and blunt, and
projects little from the succeeding whorls. The LACM-AHF collections contain ma-
terial from the Galapagos Islands, from depths of approximately 80 to 110 meters,
which seems at least tentatively referable to Dentimargo anticlea. Since these spec-
imens show evidence of maturity — e.g., slight thickening and intuming of the outer
lip and development of a low posterior denticle a short distance inside the aperture
— at lengths of 2.7 to 3.5 mm, they call into question the premise that Dali’s two
nominal taxa represent growth stages of the same species.

These specimens share with the lectotype of M. anticlea a low, blunt nuclear
shorl, a columella excavated so that the most posterior fold projects farthest into the
aperture, and an acute posterior angle of the aperture.

Referred material : Two specimens, bottom sample AHF 406, Hancock Bank,
northeast of Charles Island, Galapagos Islands, Ecuador (01° 03' 30" S, 90° 17' 30"
W), 110 m. Collected by Allan Hancock Pacific Expedition of 1933, 5 February 1933.
Five specimens, bottom sample AHF 415, north of Indefatigable Island, Galapagos
Islands (0° 27' S, 90° 22' W), 100 m. Collected by Allan Hancock Pacific Expedition
of 1933, 17 February 1933. One immature specimen, bottom sample AHF 411, north
of Duncan Island, Galapagos Islands (approx. 0° 35' S, 90° 40' W), 82 m. Collected
by Allan Hancock Pacific Expedition of 1933, 15 February 1933. Seven specimens
(four juvenile), bottom sample AHF 417, north of Indefatigable Island, Galapagos
Islands (0° 27' S, 90° 22' W), 100 m. Collected by Allan Hancock Pacific Expedition
of 1933, 17 February 1933.

A lot of three specimens, bottom sample AHF 400, Gardner Bay, Hood Island,
Galapagos Islands (approx. 01° 22' S, 89° 39' W), 31 m, collected by Allan Hancock
Pacific Expedition of 1933, 25 January 1933, is distinctive in having a medium-sized,
moderately elevated nucleus, higher and more conic spire, aperture 58% of total length
of shell (compared to an average of 66% for specimens from bottom sample AHF
415), and slightly more shouldered body whorl, resulting in a more obtuse posterior
angle of aperture. The largest specimen is 3.7 mm long. One additional, worn indi-
vidual from bottom sample AHF 407, Albemarle Point, Albemarle Island, Galapagos

1978

West American Marginellidae

13

Islands (approx. 0° 12' N, 91° 21' W), 91 m, collected by Allan Hancock Pacific
Expedition of 1933, 10 February 1933, is intermediate in spire height and projection
of the nucleus. I believe the Hood Island specimens to be extreme members in a
varying series of shallow water populations of Dentimargo anticlea.

Coan and Roth (1966: pi. 51, figs. 62, 63) illustrated as a hypotype of “Lo/-
varinella ’ eremus (Dali) a high-spired shell with orange-brown bands around the body
whorl; the same figure was reprinted by Keen (1971: fig. 1401). The figured speci-
men, from the Stanford University collection (SUPTC 9847), was labeled Albemarle
Island, Galapagos Islands. It is extremely similar to specimens of Dentimargo au-
reocincta (Stearns 1873) from Cedar Key, Florida (Locality 852, California Academy
of Sciences Department of Geology), particularly in size, disposition of the color
bands, prominence of columellar folds, and arrangement of teeth along the convex
and thickened outer lip. They also share a marked diagonal truncation of the outer lip
anteriorly. Unless further collecting in the Galapagos turns up material to confirm the
record, it appears probable that the Stanford specimen is a misallocated Atlantic shell.
The only Dentimargo species in tropical west America definitely known to have brown
banding is D. zetetes, described below.

Dentimargo zetetes new species
Figure 14

Diagnosis: A small, gracefully fusiform Dentimargo with whorls of spire only
slightly convex, sutures scarcely impressed; distinguished from other west American
species by the presence of orange-brown spiral banding which is composite in size
and strength of the bands.

Description of holotype: Shell rather small for the genus, gracefully fusiform,
somewhat inflated anteriorly. Surface highly polished, unsculptured. Translucent
white, with four narrow, sharply defined, orange brown spiral bands on body whorl,
the first band paler than the rest, just anterior to suture, the remaining three dividing
the body whorl approximately into fourths; regions between second and third, and
third and fourth bands lightly colored orange-brown, with faint indications of sec-
ondary banding; with a poorly defined orange-brown zone extending from anterior
end of shell to just anterior of last narrow band. Nucleus small, rounded, moderately
projecting. Spire high, of three whorls, flat-sided; suture indistinct, scarcely im-
pressed; two posterior color bands showing on whorls of spire. Outer lip thick, con-
vex, white with color bands continuing to its edge, with moderately broad, sloping
shoulder behind aperture; interior with one major denticle posteriorly and three
smaller, low, obscure denticles medially. Aperture moderately wide, whitish inside
with color bands showing through, anterior margin slightly produced, posterior angle
obtuse, posterior notch distinct. Parietal wall faintly convex, uncalloused. Columella
shallowly concave, with four diverging, nearly equidistant folds including fold at base
of columella; posterior three folds subequal in size, basal one smaller. Length 4. 1
mm, width 2.0 mm, length of aperture 2.2 mm.

Type locality: LACM locality 72-7 , 1-11 m, Punta Isla to 500 m south of point,

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Contributions in Science

No. 292

Bahia Elena, Guanacaste Province, Costa Rica (10° 56' 00" N, 85° 48' 55" W), col-
lected by P. LaFollette & D. Cadien, 13 February, 1972 (R/V Searcher station 382-
383). Two specimens.

Type material: Holotype, FACM 1781. Paratype, FACM 1782.

Referred material: Two specimens, AHF 472-35, Port Parker, Costa Rica (10°
57' 50" N, 85° 48' 45" W), 55 m on shell bottom. Collected by Allan Hancock Pacific
Expedition of 1935, 9 February 1935. One specimen, AHF 470-35, Port Parker, Costa
Rica (10° 57' 35" N, 85° 49' W), 9 m on sand and mud bottom. Collected by Allan
Hancock Pacific Expedition of 1935, 9 February 1935. One specimen, bottom sample
AHF 316, Gulfo Elena (Port Parker), Costa Rica (approx. 10° 55' N, 85° 49' W), 26
m. Collected by Allan Hancock Pacific Expedition of 1935, 8 February 1935. One
specimen, LACM 70-65, Playas del Coco, Costa Rica (10° 33' N, 85° 43' W), in-
tertidal. Collected by T. M. Spight, February-March 1970. Two specimens, FACM
72-30, south tip of Punta Santa Elena, Guanacaste Province, Costa Rica (10° 53' 35"
N, 85° 57' 52" W), in 12-15 m, collected by P. FaFollette and D. Cadien, 18 February
1972 (R/V Searcher sta. 415). Ninety-five specimens, FACM 72-12, 1 .6 km offshore
between Bahia Elena and Juanillo Bay, Guanacaste Province, Costa Rica (10° 57' 20"
N, 85° 46' 08" W), in 53-26 m, collected by P. FaFollette and D. Cadien, 14 February
1972 (R/V Searcher sta. 392).

Discussion: The paratype is banded similarly to the holotype, and the large lot
from FACM locality 72-12 shows minor variations of the same pattern. The banding
on a specimen from AHF 472-35 differs in having (starting below the suture) a very
faint narrow band, followed by a distinct narrow band on the posterior third of the
whorl, a broad band near the middle of the aperture, another narrow band, and a
brown zone at the anterior end of the shell. Other specimens show banding faintly
but are too worn to permit a detailed description of the pattern. The complex nature
of the banding, in both size and strength, nevertheless appears to be a diagnostic
character and helps to distinguish D. zetetes from the Caribbean Dentimargo aureo-
cincta (Steams 1873), which also differs in having a shorter aperture and denticles
borne on the edge of the outer lip rather than a short distance inside the aperture.

Dentimargo zetetes appears to be the only eastern Pacific species with a color
pattern, as mentioned above under D. anticlea. The latter species has, in most in-
stances, a larger, flatter nucleus and more convex whorls of the spire. Dentimargo
erema (Dali 1919) is unbanded, has denticles on the edge of the outer lip, and a larger,
bulbous nucleus. Dentimargo epacrodonta is unbanded, and has a narrower aperture,
narrower shoulder on body whorl, and well developed teeth inside the aperture.

Etymology: zetetes — Gr. , a searcher.

Dentimargo epacrodonta new species
Figure 15

Diagnosis: A small, ivory-white Dentimargo distinguished from other west
American species by its narrowly shouldered and anteriorly constricted body whorl
and the presence of several subequal denticles a short distance inside aperture.

1978

West American Marginellidae

15

Description of holotype: Shell rather small for the genus, narrowly biconic, pro-
duced anteriorly. Surface highly polished, unsculptured. Translucent ivory-white.
Body whorl constricted by a broad, shallow sulcus about one-third of the distance
from anterior end. Nucleus small, hemispherical, moderately projecting. Spire high,
of three convex whorls; suture appressed but distinct; “false suture” (internal trace
of each whorl’s junction with previous whorl) visible. Outer lip sharp at edge but
thickened a short distance back, with narrow, steeply sloping shoulder behind aper-
ture; straight along its medial portion; with seven distinct denticles a short distance
inside aperture, of which the most posterior and the fourth are largest, the second and
third from posterior are small and sharp, and the three most anterior are low and
progressively less distinct. Aperture moderately narrow, produced anteriorly and
somewhat effuse toward anterior end of outer lip; posterior angle acute; posterior notch
indistinct (on paratype; holotype broken here). Parietal wall excavated, not calloused.
Columella markedly concave, with four folds including fold at base of columella;
posterior two folds subparallel, anterior two very oblique. Length 3.7 mm, width 1.7
mm, length of aperture 2.0 mm.

Type locality: Bottom sample AHF 308, off Bahia Honda, Republic of Panama
(7° 44' N, 81° 35' W), 55-64 m. Collected by Allan Hancock Pacific Expedition of
1934, 21 February 1934. Three specimens.

Type material: Holotype, LACM-AHF 1783. Two paratypes, LACM-AHF 1784.

Referred material: One specimen, bottom sample AHF 312, Secas Islands, Re-
public of Panama (approx. 7° 57' N, 82° 02' W), 26 m. Collected by Allan Hancock
Pacific Expedition of 1935, 4 February 1935. Three specimens, bottom sample AHF
307, off Secas Islands (approx. 7° 50' N, 82° 00' W), 73-146 meters. Collected by
Allan Hancock Pacific Expedition of 1934, 22 February 1934.

Discussion: The ivory color, absence of banding, and narrower shoulder distin-
guish D. epacrodonta fromD. zetetes, which in addition lacks the anterior constriction
of the body whorl. No other west American species seems to have the array of aper-
tural denticles shown by D. epacrodonta, but identification should not hinge on these
characters since, as demonstrated in one paratype, the denticles may be incompletely
developed. The dentition ofD. anticlea (Dali), as far as seen, is dominated by a single
strong posterior tooth. The nucleus of D. anticlea is proportionally larger and less
projecting.

Etymology: Gr. , epakros, pointed at the end + odontos, tooth; a noun.

GENUS VOLV ARINA HINDS 1844

Type species, by subsequent designation, Redfield 1870, Marginella nitida Hinds 1844 ( =Voluta

mitrella Risso 1826); Recent, Mediterranean.

Small, cylindrical to conic; anterior notch absent; outer lip smooth or weakly
denticulate within; spire low to moderately elevated.

The status of Hyalina Schumacher 1817 (type species, H. pellucida Schumacher;
Recent, locality unknown), used by some workers for this group of species, is un-
resolved. Dodge (1955) identified//, pellucida with Voluta pallida Linnaeus 1767,
and considered the latter a recognizable Caribbean species. Woodring (1970) regarded

16

Contributions in Science

No. 292

Figures 14-16. 14. Dentimargo zetetes new species, holotype, LACM 1781, length
4.1 mm. 15. Dentimargo epacrodonta new species, holotype, LACM 1783, length
3.7 mm. 16. Volvarina innexa new species, holotype, LACM-AHF 1785, length 5.3
mm.

H. pellucida as unrecognizable and the genus as a nomen dubium. No type specimen
of H. pellucida is extant (J. Knudsen, personal communication); designation of a
neotype in conjunction with a revisionary study would resolve the question.3

Volvarina innexa new species
Figure 16

Diagnosis: A small Volvarina with relatively high spire, conic outline, and large
nucleus, differing from other west American species by being white with two trans-
lucent spiral bands and having a narrow yellow band near the anterior end.

Description of holotype: Shell small for genus, elongate-conic, narrower ante-
riorly. Surface highly polished, unsculptured. Translucent white with two spiral zones
of greater translucency, the first extending from shoulder of body whorl to just below
suture, the second at middle of body whorl. Outer lip and extreme anterior end opaque
white. Nucleus large; spire elevated, nearly transparent, with “false suture” (internal
trace of each whorl’s junction with previous whorl) visible; whorls of spire convex.
Outer lip varicose externally, thickest posterior to middle, slightly constricted medi-
ally, not denticulate. Aperture narrow, evenly widening to anterior end; anterior mar-

3Coan and Roth (1976, Jour. Moll. Stud. 42:217-222) recently discussed this problem and des-
ignated a neotype.

1978

West American Marginellidae

17

gin evenly rounded, without indentation; posterior notch shallow. Parietal wall mod-
erately convex, uncalloused. Columella very slightly concave, with four equidistant,
nearly parallel folds including fold at base of columella, the folds terminating distally
in a weakly defined white fasciole with a pale yellow spiral band along its posterior
margin. Length 5.3 mm, width 2.2 mm.

Type locality: Station AHF 177-34, Sullivan Bay, James Island, Galapagos Is-
lands, Ecuador (0° 16' 30" S, 90° 35' 15" W), 9-37 m on bottom of rock with sand
patches. Collected by Allan Hancock Pacific Expedition of 1934, 23 January 1934.
One specimen.

Type material: Holotype, L ACM- AHF 1785.

Discussion: The high spire, prominent nucleus, and conic outline of Volvarina
innexa distinguish it from other west American species of the genus, as does its sin-
gular color pattern. Volvarina nyssa Roth and Coan 1971, from Pinta and Genovesa
Islands, Galapagos Islands, has a much lower spire and bright orange-brown bands
on the body whorl. Volvarina taeniolata rosa (Schwengel 1938) is larger, more ovate
in outline, and bright pink in color; some specimens show brown banding like that
of the nominate subspecies, which ranges from southern California to Central Amer-
ica. Some specimens from the Gulf of California, like those referred to Volvarina
taeniolata by Coan and Roth (1966: pi. 50, figs. 54, 55), resemble V. innexa in their
pale coloration and high spire. They do not however have the large nucleus of the
new species. The columellar folds of V. innexa seem to be proportionally the largest
of any west American Volvarina examined. I interpret V. innexa as an eastern Pacific
representative of the group of high-spired Caribbean Marginellidae which includes
Volvarina veliei (Pilsbry 1896) and V. avenacea (Deshayes 1844).

Etymology: innexa — L. , joined.

RESUMEN

Se describen seis nuevas especies de Marginellidae: Prunum ( Prunum ) gorgo-
nense de la Isla Gorgona, Colombia; Prunum {Prunum) macleani de la Republica del
Ecuador; Prunum ( Microspira ) aletes y Dentimargo zetetes de la costa Pacifica de
Costa Rica; Dentimargo epacrodonta de la Republica de Panama; y Volvarina innexa
de las Islas Galapagos. Prunum ( Microspira ) aletes se parece a algunas especies Car-
ibes y a especies Neogenos de Florida. Prunum {Prunum) curtum (Sowerby) se ex-
tiende hasta aguas templados al sur de la provincia Panamica; su presencia en el Golfo
de Panama requiere confirmacion. La radula de Prunum {Prunum) woodbridgei (Her-
tlein y Strong) se ilustra; concuerda bien con la radula de Prunum storerium (Couth-
ouy). Dentimargo anticlea (Dali) — identificacion tentativa — se encuentra en varias
localidades en las Islas Galapagos.

LITERATURE CITED

Abbott, R. T. 1957. Prunum roscidum in New Jersey. Nautilus 7(2):52-53, pi. 4, figs. 4, 4a.
. 1958. The marine mollusks of Grand Cayman Island, British West Indies. Acad. Nat.

Sci. Philadelphia, Monogr. 11:1-138, pis. 1-5.

18

Contributions in Science

No. 292

Akers, W. H.1974. Age of Pinecrest Beds, south Florida. Tulane Stud. Geol., Paleontol.

1 1(2): 1 19-120.

Bayer, F. M.1943. Observations on marine Mollusca, with descriptions of new species. Nau-
tilus 56(4): 109-1 15, pis. 12, 14.

Carcelles, A. 1944. Catalogo de los moluscos marinos de Puerto Quequen (Republica Argen-
tina). Revista del Museo de La Plata (N.S.) 3 (Zool. 23):233-309, pis. 1-15.

Clench, W. J. and R. D. Turner. 1950. The western Atlantic marine mollusks described by
C. B. Adams. Occas. Pap. Moll. l(15):233-403, pis. 29-49.

Coan, E. V. and B. Roth. 1966. The west American Marginellidae. Veliger 8(4):276-299,
pis. 48-51, text-figs. 1-5.

Dall, W. H. 1890. Contributions to the Tertiary fauna of Florida, with especial reference to the
Miocene silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Part I.
Pulmonate, opisthobranchiate and orthodont gastropods. Wagner Free Inst. Sci. Philadel-
phia, Trans. 3(l):l-200, pis. 1-12.

. 1892. Contributions to the Tertiary fauna of Florida, with especial reference to the

Miocene silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Part II.
Streptodont and other gastropods, concluded. Wagner Free Inst. Sci. Philadelphia, Trans.
3(2):20 1 -473 , pis. 13-22.

. 1919. Descriptions of new species of Mollusca from the North Pacific Ocean in the

collection of the United States National Museum. U. S. Nat. Mus., Proc. 56(2295):293-
371.

Dodge, H.1955. A historical review of the mollusks of Linnaeus. Part 3. The genera Bulla and
Voluta of the class Gastropoda. Amer. Mus. Nat. Hist., Bull. 107(1): 1-158.

Hertlein, L. G. and A. M. Strong. 1951. Descriptions of three new species of marine gas-
tropods from west Mexico and Guatemala. So. California Acad. Sci., Bull. 50(2):76-80,
pi. 26, figs. 1-4, 8.

Keen, A. M.1971. Sea shells of tropical west America: Marine mollusks from Baja California
to Peru. Stanford Univ. Press, Stanford, California, xiv + 1064 p., 22 pis. [Marginellidae,
pp. 632-638, by Coan and Roth.]

Olsson, A. A. 1964. Neogene mollusks from northwestern Ecuador. Paleontol. Res. Inst., Ith-
aca, New York, 256 p., 36 pis.

. 1967. Some Tertiary mollusks from south Florida and the Caribbean. Paleontol. Res.

Inst., Ithaca, New York, 61 p. , 8 pis.

Reeve, L. A. 1864 [1864-1865], Monograph of the genus Marginella. In: Conchologia iconica:
or illustrations of the shells of molluscous animals 15, pis. 2-13 [1864], pis. 1, 14-27,
index, errata [1865].

Roth, B. and E. V. Coan. 1971 . Marginellidae (Mollusca: Neogastropoda) from the Galapagos
Islands and Cocos Island. Proc. Calif. Acad. Sci. (4)37(23):575-584, text-figs. 1-5.

Sowerby, G. B., 1.1832. Characters of new species of Mollusca and Conchifera collected by
Mr. Cuming. Proc. Zool. Soc. London 2 [part 2] (18): 104- 108.

Tryon, G. W., Jr. 1 882[ 1 882- 1 883] . Family Marginellidae. In: Manual of conchology; struc-
tural and systematic 5. Marginellidae (17): 5-58, pis. 2-13 [1882]; (20):197-255 [1883].

Woodring, W. P.1970. Geology and paleontology of Canal Zone and adjoining parts of Pan-
ama. Description of Tertiary mollusks (Gastropods: Eulimidae, Marginellidae to Helminth-
oglyptidae). U. S. Geol. Surv., Prof. Paper 306-D:299-452, pis. 48-66.

Accepted for publication October 27, 1976.

5 Ol> 7

0mmw^

MAY 2 2 1978 *

NUMBER 293
MAY 15, 1978

MCCOSKERICHTHYS SANDAE,
A NEW AND UNUSUAL CHAENOPSID BLENNY
FROM THE PACIFIC COAST OF PANAMA AND COSTA RICA

By Richard H. Rosenblatt and John S. Stephens, Jr.

■

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

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MCCOSKERICHTHYS SANDAE, A NEW AND UNUSUAL CHAENOPSID

BLENNY

FROM THE PACIFIC COAST OF PANAMA AND COSTA RICA1’2

by Richard H. Rosenblatt3 and John S. Stephens, Jr.4

Abstract: A new genus and species of chaenopsid blenny is described, based
upon a series of 651 specimens. This new species occurs at depths ranging from
1 to 30 meters along the Pacific coast of Costa Rica and Panama. It is a tube dwelling
biennioid fish, living along near- vertical rock faces or coral heads. This new chaenop-
sid is unique in having the following combination of characters: four circumorbital
bones, ectopterygoid teeth, jaw with spatulate teeth on outer row but granular ones
on inner row, five orbital cirri, a blunt head, 17-20 spines and 31-34 soft rays in
dorsal fin, 2 spines and 32-36 soft rays in anal fin, and 54-57 vertebrae. The rela-
tionships of the new genus are discussed as they relate to the clinidlike blenny families
(Clinidae, Chaenopsidae, Tripterygiidae). Osteological features are discussed and il-
lustrated, and their interpretation as evidence of evolutionary significance is presented.

INTRODUCTION

In 1971 while on fellowship to the Smithsonian Tropical Research Institute in
Panama, John E. McCosker collected and made observations on a new species of tube
dwelling biennioid fish. Extensive series of this species were also taken by McCosker
and Rosenblatt during the Janss Foundation expedition to Costa Rica on the R/V
SEARCHER . The characteristics of this species are so distinctive that it cannot be
assigned to any known genus, but its characteristics certainly ally it to the family
Chaenopsidae.

’Review Committee for this Contribution
Robert J. Lavenberg
Richard Robins
Victor G. Springer

Contribution from the Scripps Institution of Oceanography.

3Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California
92093.

4 Department of Biology, Occidental College, Los Angeles, California 90041.

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Mccoskerichthys new genus
Figure 1

Diagnosis: A chaenopsid with the following unique combination of characters:
four circumorbital bones, teeth on the ectoptergoid, outer row of jaw teeth spatulate,
teeth of inner rows granular, 5 pairs of orbital cirri and a combination of blunt head
and high dorsal, anal and vertebral counts. Other characters those of the single in-
cluded species.

Type species: Mccoskerichthys sandae new species

Derivation of name: Named for John E. McCosker, who discovered the species
and has participated in the collection of all known specimens.

RELATIONSHIPS

Bohlke and C. H. Robins (1974) in describing the new clinid genus Haptoclinus
discussed certain characters of this monotypic genus that they regarded as breaking
down the separation of the three often recognized clinidlike blenny families: the Clin-
idae, Chaenopsidae and Tripterygiidae. They recognized only the family Clinidae,
admitting that the chaenopsids and tripterygiids represent lines of divergence within
the family, but suggested that other equally distinct clinid subgroups may later be
defined. However, the characters given for Haptoclinus apectolophus do not seem to
indicate relationship to chaenopsids; in fact, all its characters are typically clinid.
There is also very little evidence presented allying Haptoclinus to the tripterygiids.
Haptoclinus represents a specialized clinid, perhaps allied to Paraclinus .

It may well be that uniting all clinidlike or clinid-derived fishes into a single
family is the best way to treat these relationships but unfortunately the discussion by
Bohlke and Robins (1974) is mostly irrelevant to this problem. We, therefore, will
continue to treat clinids, chaenopsids and tripterygiids as separate blennioid families.

The morphological characters given in the diagnosis of Mccoskerichthys indicate
its isolated position within the family Chaenopsidae. Any one of these characters dis-
tinguishes this genus from all other known chaenopsids, and the presence of ectop-
teryoid teeth is unique within the Blennioidei. If these special characters are ignored,
Mccoskerichthys seems to be most closely related to the Protemblemaria to Acan-
themblemaria line of chaenopsids in that the paired nasals have fused medially into

Figure 1 . Mccoskerichthys sandae, Holotype

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New Chaenopsid Blenny

3

a single element and the frontals (parietal of Jollie 1962; Stephens 1963) are ridged
and sculptured (Fig. 3). Further, there is a tendency toward modified dentition in this
evolutionary line. The species of Chaenopsis agree with Mccoskerichthys in having
a high number of vertebrae and dorsal and anal soft rays, as well as the correlated
characters of a reduced number of caudal peduncle vertebrae and the absence of dorsal
pterygiophores anterior to the first vertebra, but are otherwise very different.

Stephens (1963) listed 13 characters he considered to represent the primitive con-
dition in chaenopsids. Mccoskerichthys exhibits the primitive condition in six char-
acters (snout shape, orbital cirri, palatovomerine dentition, upper jaw length dimor-
phism, median fin height and circumorbital width). Although relatively slender
circumorbitals were considered to be primitive by Stephens, this character needs clar-
ification. The primitive condition is better described as a slender lachrymal and the
advanced character a ventrally expanded lachrymal as in most Coralliozetus (8 8 )and
most species of Acanthemblemaria (Figs. 6-9). The other circumorbitals are relatively
stout primitively (Stephens 1963, fig. 2A and B). The slender circumorbital ring is
found in almost all species of the Emblemariopsis to Chaenopsis line, though several
species of Emblemaria show a modified, stouter ring. The circumorbital ring of
Mccoskerichthys is similar to that of Neoclinus, a clinid considered close to the an-
cestral chaenopsid lineage, in that there are four bones, the lachrymal is slender and

Figure 2. Mccoskerichthys sandae, a living individual in a tube in the coral panona.
Photo taken by Edwin Janss at Isla del Cano Costa Rica in March, 1973.

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the other three circumorbitals (postorbitals) relatively stout. Neoclinus has a distinct
dermosphenotic not present in Mccoskerichthys or other chaenopsids.

The four circumorbitals, blunt snout and low nondimorphic dorsal fin impart a
primitive facies to Mccoskerichthys . The strongly modified dentition, presence of teeth
on the ectopterygoid, and the elongate body are specializations but do not argue
against a primitive origin. If the presence of four circumorbital bones indeed represents
the retention of a primitive character, then Mccoskerichthys must have evolved prior
to the division of the chaenopsids into the Protemblemaria to Acanthemblemaria and
Emblemariopsis to Chaenopsis lineages, as the circumorbitals are reduced in both.
If Mccoskerichthys diverged this early, however, it is difficult to explain its numerous
apomorphic characters shared with members of the Protemblemaria to Acanthemble-
maria lineage, i.e. a single fused nasal, cranial sculpturing, specialized incisor and
molariform dentition, as well as short and stout pelvic fins.

Mccoskerichthys is certainly not particularly clinidlike: the body is scaleless, and
the lateral line absent, most of the head pores are simple, there is a single epural, and
the fifth hypural (minimal hypural) is not identifiable nor are the dermosphenotic and
median and lateral extrascapulars. All of these are chaenopsid characters. Typically
there are two epurals and the fifth (minimal) hypural in the Clinidae and the der-
mosphenotic and one or more extrascapulars are present. It must be stressed, however,
that the chaenopsid condition in these osteological characters is approximated by
Stathmonotus and Starksia and in one or more characters by other clinids. An addi-
tional chaenopsid character, the presence of a well defined anteriorly projecting lateral
process on the hyomandibular (Figs. 6, 7, and 9) has been examined by Springer
(personal communication). This hook is certainly characteristic of most species but
appears to be absent in Coralliozetus (Fig. 8) and those species of Emblemariopsis
that we have examined. This process is at best rudimentary in Mccoskerichthys (Fig.
3).

It is difficult to evaluate the phyletic significance of the presence of four separate
postorbital bones in Mccoskerichthys . It would be especially helpful to know whether
there are three postorbital centers of ossification or only one in the rest of the chaen-
opsids. If there are three, joints could easily be regained; if one, it is likely that the
separate bones of Mccoskerichthys represent a primitive condition. The fact that the
postorbital of other chaenopsids anomalously may show separate elements suggests
that several centers of ossification are present and that these normally fuse during
development.

It is our interpretation, based on parsimony, that Mccoskerichthys evolved after
the main chaenopsid features had appeared, and that it represents an offshoot of the
Protemblemaria to Acanthemblemaria line close to Ekemblemaria in the phylogenetic
diagram given by Stephens (1970) but has lost the fusion of the postorbitals. This
phyletic position eliminates the difficulty of interpreting the synapomorphies of the
Mccoskerichthys and the Protemblemaria to Acanthemblemaria specjes.

Mccoskerichthys also agrees with the Chaenopsidae in another feature, which has
not previously been noted. In chaenopsids the maxilla is not visible externally. An-
teriorly it slips beneath the lower free margin and posterior projection of the lachrymal
(the postorbital or second circumorbital usually forms the dorsal parts of the projec-

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5

tion). Posteriorly, a free fold of skin extends from the postero ventral angle of the
circumorbitals across the cheek forming a pocket covering the maxilla. This pocket
is variously developed but present in all species. It is best developed in Chaenopsis ,
in which the maxilla extends far beyond the rear margin of the eye, and least developed
in forms where the maxilla is short or the posterior expansion of the circumorbital
ring is large. In Mccoskerichthys the posterior extension of the lachrymal and the
second circumorbital cover all but the posterior tip of the maxilla and the free fold
is rudimentary. Emblemaria has a reduced fold, but the maxilla is buried beneath the
skin and there is a fold across its tip. This character is approximated in some genera
of clinids including Stcithmonotus and Malacoctenus but not Haptoclinus .

The effect of these structural chaenopsid modifications is to limit the lateral ex-
pansion of the jaws. Because of the limited protusibility of the premaxilla, the maxilla
is but little rotated downward when the mouth is opened. Illustrations of threat display
in which the mouth is opened widely in Chaenopsis and Emblemaria (Robins, Phillips
and Phillips 1959, text fig. 1, PI. 3, figs. 1 and 2; Wickler 1964) indicate that the
maxilla remains closely applied to the cheek. The rigidly fused circumorbital ring of
chaenopsids further limits lateral mobility of the maxilla.

It is possible that the early chaenopsids were omnivorous nibblers, and that the
change to plankton-picking, characteristic of most members of the family (species of
Coralliozetus , Acanthemblemaris , Protemblemaria, and Emblemaria examined, see
also Stephens, Hobson and Johnson 1966; Hobson 1968) was correlated with increas-
ing reliance on the tubiculous habit. Chaenopsis and Lucayablennius (Hobson 1968;
Greenfield 1972) have become predators on relatively large food items, and the gape
is proportionately long, but the basic chaenopsid jaw structure is otherwise little mod-
ified. The chisellike teeth of Mccoskerichthys hardly seem adapted to plankton feed-
ing, yet observations and gut analyses suggest this mode of existence.

Mccoskerichthys sandae new species
Figures 1, 2

Morphology: Terminology is that of Stephens 1970. Counts and measurements
of certain body parts are given in Tables 1 and 2. Body greatly elongated for a chae-
nopsid, approaching the species of Chaenopsis in this respect, body depth at dorsal
origin 10-12 in standard length. Body strongly compressed; its dorsal and ventral
outlines with little posterior taper. Head length 5. 5-7.0 in standard length, head width
slightly less than its depth. Snout short and rounded, lips projecting because of broad,
projecting incisors; snout length from two-thirds of, to almost equal to, eye diameter,
difficult to measure accurately, because of anterior excavation of orbit. Eyes directed
anteriad as well as laterad, pupil round; diameter of eye 3. 5-4. 5 in head. Interorbital
narrow, rim of orbit incised anteriorly to accommodate anteriorly directed eyes, and
posteriorly to accommodate orbital cirri. Orbital cirri forming a bushy mass, which
completely covers interorbital region. Cirral mass composed of five pairs of cirri; each
cirrus with a stout base and at least two major branches, one medial and one lateral,
each branch complexly multifid and with short lateral branches, imparting a ragged

6 Contributions in Science No. 293

Table 1

Meristic variations in

Mccoskerichthys sandae.

Total Dorsal Elements

50

51

52

53

8

19

14

6

Dorsal Spines

XVII

XVIII

XIX

XX

4

12

26

5

Dorsal Soft Rays

31

32

33

34

3

17

19

8

Anal Soft Rays

32

33

34

35 36

1

1

23

21 1

Pectoral Rays*

12

13

14

5

85

2

Total Vertebrae

54

55

56

57

6

32

12

4

*2 damaged pectoral fins with 10 and 1 1 rays

appearance. Cirral mass extends from posterior edge of postorbital flange of frontal
to anterior margin of eye at level of center of pupil; height of mass slightly less than
eye diameter.

Top of head behind eyes with a series of transverse folds, which become lon-
gitudinal and converge toward dorsal commissural pore; folds originate anteriorly
about at junction of frontals with circumorbitals and reflect underlying bony ridges.
A crescentic fold at occiput, outlining anterior extent of body musculature. Fold re-
duced in females and absent in juveniles, although bony ridges may be seen.

Upper jaw horizontal, terminating behind eye, its length about 1.3-1. 5 in post-
orbital head length. Maxilla in males extending slightly beyond end of posteroventral
expansion of circumorbitals, rather than ending beneath it as in females. Upper jaw
with a pair of recurved incisors, one on either side of symphysis, followed by three
enlarged, somewhat forward-directed, spatulate incisors; posterolateral to these a row

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Table 2

Measurements in millimeters of body parts of the holotype and
ten paratypes of Mccoskerichthys sandae

Holotype

Standard length

62.5

63.5

65.0

69.5

Head Length

10.0

9.6

10.0

10.0

Head Depth

5.5

5.1

5.6

5.9

Head Width

5.4

5.3

5.7

5.5

Upper Jaw Length

4.9

4.3

4.5

4.5

Snout Length

1.9

1.7

1.9

2.0

Eye Diameter

2.1

2.3

2.1

2.0

Interorbit Width

.9

.9

.9

.9

Predorsal Length

8.0

7.9

7.9

8.5

Preanal Length

25.5

25.0

26.1

27.8

Body Depth at Anus

5.7

4.8

5.3

5.2

62.6

59.9

62.5

62.0

69.1

65.7

65.0

9.7

9.9

9.6

9.6

9.3

9.6

9.5

5.0

4.9

5.5

5.1

5.6

5.4

5.5

5.0

5.1

5.9

5.4

5.8

5.4

5.1

4.5

4.9

4.9

4.0

5.0

4.6

4.4

1.6

1.6

1.9

1.9

1.9

1.9

1.5

2.0

2.3

2.1

2.1

2.0

2.3

2.1

.8

.7

.7

.6

.8

.6

.7

7.9

8.0

8.0

7.9

8.0

8.0

7.8

24.6

24.5

25.0

25.4

26.0

27.0

25.6

4.9

4.8

5.5

5.6

5.6

5.0

5.5

of 7-10 smaller, downward directed, stout incisors. A patch of granular teeth ante-
riorly behind large incisors, continued as a single row behind small incisors. Dentition
of lower jaw similar to that of upper except that there are no small symphysial incisors,
and there is a pair of stout, conical, almost caniniform teeth, one on each side of
symphysis, behind patch of granular teeth. About 6 short, stout, teeth on vomer. Palate
with a patch of teeth anteriorly on palatine, narrowing to two rows posteriorly; ec-
topterygoid with a zig-zag row of ten teeth. Anterior nostril just above upper lip, with
a short tube bearing two tentacles on its rim, one anterior and one posterior. Posterior
nostril just behind anterior. Cranial sensory pores mostly simple, except over postor-
bital bones; preoperculomandibular series with 1 commissural, 3 occipital, 1 lateral,
and 1 temporal, 6-7 preopercular and 5 mandibular anterior to corner of mouth; post-
orbital series with 2 pores; supraorbital series with 4 pores. No pores on body posterior
to upper corner of gill opening.

Gill membranes united, free from isthmus; branchiostegals 6. Gill rakers 4-
5+11-12 on first arch in five specimens counted.

Total dorsal fin rays 50-53; XVII-XX, 31-34. Dorsal origin above upper anterior
comer (formed by attachment of opercle membrane to body) of operculum. All rays
evenly spaced, first spine short, two-thirds length of second, subsequent rays sub-
equal, with middle of soft dorsal somewhat higher; longest soft rays 1.5 X first and
last soft rays.

No notch between spinous and soft dorsal, last ray attached for its entire length
by a membrane inserting on middle third of dorsalmost segmented caudal ray. Anal
fin II, 32-36, all soft rays evenly spaced, spines slightly closer together. First spine
short, one-half length of second, and one-third as long as first soft-ray. Soft-rays
subequal, last ray attached as last dorsal ray but to ventralmost procurrent caudal ray.
Caudal rounded, with 13 segmented rays and 3-6 procurrent rays; only one procurrent
ray visible dorsally and one ventrally. Pectorals rounded, about 1.5 in head length,
rays 12-14. Pel vies short, about two-thirds pectoral length, I, 3, spine short, closely

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applied to first soft-ray, soft-rays joined by membrane, middle ray longest. Male with
a well-developed, fingerlike genital papilla, female with numerous short, flattened,
fleshy processes around anus, and a short, broad genital papilla.

Osteological characteristics: The important cranial and caudal characters of
Mccoskerichthys sandae are presented in figures 3-5. Comparative illustrations of
Protemblemaria to Acanthemblemaria are presented in figures 6-14. Worthy of note
for Mccoskerichthys are: the specialized anterior dentition, well-hidden maxillary,
posteroventrally expanded circumorbital ring; 3 postorbital elements; fused nasals,
sculptured frontals; single epural oriented almost vertically; absence of distinct min-
imal hypural; close association of last dorsal and anal pterygiophore to caudal (3 ver-
tebrae between urostyle and last pterygiophore). Two additional osteological char-
acteristics of this species that may be worthy of note are the presence of 3 anal
pterygiophores anterior to the first haemal arch (typical of chaenopsids) and the lack
of any dorsal pterygiophores anterior to the first vertebra (all chaenopsids except
Chaenopsis have 2-3 pterygiophores anterior to the first vertebra).

Color in alcohol: Males are dark brown to tan with a series of darker marks on
the sides. The dark markings vary from squarish or almost circular blotches to bars.
When blotches are present there is usually a series of about 8 along the dorsal base
and a series of 10-12 more or less centered on the midline. When bars are present
there are usually about 10-12. The bars are irregular, and may be split by light centers.
The head is notably lighter than the body, but there is much variability. In some
individuals only the snout, jaws and lower surface of the head are much lighter than
the sides. The cheeks and opercles are usually darker than the anterior and ventral
parts of the head. The orbital cirri appear pale, but are dusted with melanophores.
Often the light coloration of the head extends onto the body and there is a sharp line
of demarcation extending from the pectoral base to the dorsal, between the third and
fourth spines. The line of demarcation is accentuated by a dark bar behind the fourth
dorsal spine. The belly is dusted with chromatophores and varies from lighter than
the sides to almost as dark as them. The folds around the anus are outlined in black
and the genital papilla is contrastingly white. There is much variability in coloration:
in some individuals the body is light with the dark markings strongly contrasted, in
others the body is dark brown and the markings difficult to discern. This variation in
coloration is not correlated with size, but may be related to breeding condition.

The median fins are clear, except that the dorsal fin is marked with brown for
the first 3-5 spines. The pectoral has a dusky area basally and the pelvics usually have
a few melanophores on them, mostly basally.

Females are similar to males in coloration, but the head, orbital cirri and belly
are paler, and a higher proportion have pale background color. However, dark females
and light males do occur. In life the impression is of a dark red fish with the light
areas light olive green. The dark body markings are vague, and all the fins are red.

Ecology: M. sandae occurs along near- vertical rock faces or coral heads, in areas
of moderate surge, where they inhabit mollusc or worm tubes. The heads protrude
from the tubes, and the fish make short darts forward, apparently snapping at plankton.
Individuals were not seen to emerge completely from their tubes, but may come out

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New Chaenopsid Blenny

9

for one-half the body length or more. The elongated body of M. sandae allows the
head to be thrust well away from the tube in feeding, with the tail still in contact with
the substrate. Stephens et al. (1966) indicated that Acanthemblemaria macrospilus ,
a relatively short bodied species, may leave its tube in feeding.

Observations indicating thatM. sandae feeds on plankton were confirmed by the
examination of gut contents of several specimens. Most individuals contained frag-
ments of copepods and amphipods, and occasional ostracods. Individuals from LACM
32551 contained numbers of tiny (.3-. 5 mm) planktonic stages of pelecypods and
gastropods.

M. sandae has been taken at depths ranging from 1m (3 ft) to almost 30 meters
(100 ft). The shallowest records were from the Gulf of Chiriqui, Panama; at Isla del
Cano, Costa Rica; individuals were not seen much shallower than 5 meters (15 ft).

There was a striking disparity in the numbers of males and females in the Costa
Rica collections SIO75-404 (capture depth 5-10 m) contained 149 males and 76
females. LACM 32551 (capture depth 20-30 m) yielded 231 males and 154 females.
The ratios of males to females deviate from 50:50 at p < .01. This unbalanced sex
ratio in favor of males is difficult to explain. It almost certainly does not represent
unisexual patchiness. The areas collected were large, and the tube-dwelling habit,
small size and relatively poor swimming ability of the species makes it unlikely that
the sexes could come together for breeding if they did not live in close proximity. It
is possible that collecting bias toward the more obvious specimens could be involved,
since males reach a larger size than females. This cannot be ruled out, but it is not
likely that the great disparity in sex ratios can be attributed to it.

Otoliths were removed from three specimens before preservation (63.0, 66.0 and
69.5 mm). The otoliths were cleared in anise oil and examined by John E. McCosker.
The largest and smallest specimens had three hyaline rings, the outer at the margin,
indicating that they may have been entering their fourth year of growth. The 66.0 mm
individual did not have differentiated areas in the otolith.

Range: Thus far known only from Isla del Cano, Costa Rica and the Golfo de
Chiriqui, Panama.

Derivation of name: Named for Sandra McCosker, who participated in the first
collection of the species.

Material Examined

Holo type: LACM 32551-34, a 62.5 mm 6 from an isolated rockpile 1.4 miles
NW of Isla del Cano, Costa Rica. Taken with rotenone in 65' -75' (21-24 m) by
William Bussing, John McCosker, James McLean, Manuel Murillo, Richard Rosen-
blatt and Richard Wheeler on 16 March 1972.

Paratypes: Costa Rica: LACM 32551-35, 403 (18-70 mm), bearing the same
data as the holotype; USNM 214706, 10, bearing the same data as holotype; SIO-
75-404, 255 (21.5-62), Isla del Cano. Panama: Golfo de Chiriqui; SIO70-358, 14
(22-65 mm), Isla Montuosa; SIG71-51, 6 (30-60 mm), SI071-37, 2 (42-46 mm), Isla
Uva; SI071-54, 1 (38), Isla Canal de Afuera.

PRO

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Figure 3. Cranial skeleton, Mccoskerichthys sandae (SIO70-358)

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New Chaenopsid Blenny

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PTO

M. SANDAE

Figure 4. Neurocranium, Mccoskerichthys sandae (SIO70-358)

DORSAL PTERYGIOPHORE

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<i

q:

<

<

M. SANDAE

CAUDAL

Caudal skeleton, Mccoskerichthys sandae (SIO70-358)

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New Chaenopsid Blenny

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Figure 6. Cranial skeleton, Protemblemaria bicirris (UCLA 65-71)

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Figure 7. Cranial skeleton, Ekemblemaria myersi (UCLA 53-86)

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New Chaenopsid Blenny

15

>-

<

o

LU

o

z

<

o

Figure 8. Cranial skeleton, Coralliozetus angelica (right side damanged) (UCLA 65-86)

SHIV,

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Figure 9. Cranial skeleton, Acanthemblemaria castroi (UCLA 64-33)

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New Chaenopsid Blenny

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P BICIRRIS

Figure 10. Neurocranium of Protemblemaria bicirris (UCLA 65-71)

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E MYERSI

Figure 1 1. Neurocranium of Ekemblemaria myersi (UCLA 53-86)

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New Chaenopsid Blenny

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C ANGEUCA

Figure 12. Neurocranium of Coralliozelus angelica (UCLA 65-86)

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New Chaenopsid Blenny

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CAUDALS

Figure 14. Comparative caudal skeletal anatomy, Protemblemaria to Acanthemble-
maria line.

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ACKNOWLEDGMENTS

The Costa Rica collections were made possible by the Janss Foundation; the
Panama collections were made from the R/V ALPHA HELIX.

The illustration of the holotype was drawn by Brian Burnette. The osteological
drawings are by Stephens except for figure 6 by Cassie Cussik. The osteological por-
tion of this study was partially supported by National Science Foundation Grant
GB27266 to Stephens.

A NOTE ABOUT FIGURES AND LEGENDS

The osteological figures do not attempt to illustrate all sensory pores. The fol-
lowing abbreviations are used in the osteological illustrations.

AN, Angular; AR, Articular; BOC, Basioccipital; BR, Branchiostegals; BS,
Basisphenoid; CO, Circumorbital; D, Dentary; ECT, Ectopterygoid; EPO, Epiotic;
EXO, Exoccipital; F, Frontal; HYM, Hyomandibular; IC, Intercalar; IOP, In-
teropercular; L. Lachrymal; LE, Lateral ethmoid; MPT, Metapterygoid; MSPT,
Mesopterygoid; MX, Maxilla; N, Nasal; OP, Opercular; P, Parietal; PAL, Pal-
atine; PMX, Premaxilla; POP, Preopercular; PRO, Prootic; PS, Parasphenoid;
PTO, Pterotic; PTS, Pterosphenoid; Q, Quadrate; SOC, Supraoccipital; SOP,
Subopercular; SPO, Sphenotic; SYM, Symplectic; V, Vomer.

LITERATURE CITED

Bohlke, J. E., and C. H. Robins. 1974. Description of a new genus and species of clinid fish
from the western Caribbean, with comments on the families of the Blennioidea. Proc. Acad.
Nat. Sci. Phil. 126, 1:1-8.

Greenfield, D. W. 1972. Notes on the biology of the arrow blenny, Lucayablennius zingaro
(Bohlke) from British Honduras. Copeia 1972, (3):591-2.

Hobson, E. S. 1968. Predatory behavior of some shore fishes in the Gulf of California. U. S.
Dept. Interior, Bureau Sport Fish, and Wildlife, Res. Report 73:1-92.

Jollie, M. 1962. Chordate Morphology. Reinhold Publ. Corp., N. Y., 478 p.

Robins, C. R., C. Phillips and F. Phillips. 1959. Some aspects of the behavior of the blen-
nioid fish Chaenopsis ocellata Poey. Zoologica, 44, pt. 2 (5):77-83.

Stephens, J. S., Jr. 1963. A revised classification of the blennioid fishes of the American
family Chaenopsidae. Univ. Calif. Publ. Zook, 68:1-165.

— . 1970. Seven new chaenopsid blennies from the western Atlantic. Copeia 1970 (2):

280-309.

Stephens, J. S., Jr., E. S. Hobson and R. K. Johnson. 1966. Notes on distribution, behavior,
and morphological variation in some chaenopsid fishes from the tropical eastern Pacific,
with descriptions of two new species, Acanthemblemaria castroi and Coralliozetus spring-
eri. Copeia 1966 (3):424-438.

Wickler, W. 1964. Emblemaria pandionis (Blenniidae). Kampfverhalten. Encyclopedia Ci-
nematographica E 5171, 1963. 3 p.

Accepted for publication June 30, 1976.

ZL%&9>

NUMBER 294
MAY 15, 1978

A NEW SPECIES OF CHIROPTEROTRITON (AMPHIBIA: CAUDATA)
FROM BAJA VERAPAZ, GUATEMALA, WITH COMMENTS ON
RELATIONSHIPS AMONG CENTRAL AMERICAN
MEMBERS OF THE GENUS

By James F. Lynch and David B. Wake

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCENCC

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A NEW SPECIES OF CHIROPTEROTRITON (AMPHIBIA: CAUDATA) FROM
BAJA VERAPAZ, GUATEMALA, WITH COMMENTS ON RELATIONSHIPS
AMONG CENTRAL AMERICAN MEMBERS OF THE GENUS1

By James F. Lynch2 and David B. Wake3

Abstract: Chiropterotriton veraepacis, a diminutive species of plethodontid
salamander with very large nostrils, is described. It occurs inside bromeliads in an
area of montane cloud forest south of Purulha, Baja Verapaz, Guatemala, and in
nearby areas of Alta Verapaz, Guatemala. In most structural features C. veraepacis
resembles C. nasalis of northeastern Honduras, a shorter tailed, wider headed species
with larger nostrils. The new species is similar to species of the bromeliacia group
from Guatemala and Chiapas in ecology and in many structural features but differs
from them in osteology. Although C. nasalis and C. veraepacis are very distinct from
the Costa Rican species C. picadoi in most morphological and ecological features,
they resemble that species in osteology. Some additional information is presented
concerning the poorly known species C. richardi (Costa Rica) andC. barbouri (Hon-
duras), and possible relationships among all of these species are discussed.

INTRODUCTION

In a recent paper (Lynch and Wake 1975) we assigned two new Guatemalan
species of Chiropterotriton (C. cuchumatanus and C. rabbi ) as well as three previ-
ously known Chiapan and Guatemalan forms (C. megarhinus, C. xolocalcae, and C.
bromeliacia ) to the bromeliacia species group. In that paper we noted the importance
of Nuclear Central America as an evolutionary center for Chiropterotriton and other
tropical plethodontid genera (see also, Wake and Lynch 1976), and remarked that
other undescribed Chiropterotriton existed in the area.

This paper reports the discovery of a new species of Chiropterotriton from the
Department of Baja Verapaz, in east-central Guatemala. This form was first encoun-
tered in the spring of 1973 by Thomas Uzzell, Academy of Natural Sciences, Phil-
adelphia (ANSP), who collected four individuals in an area of montane cloud forest
a few kilometers south of the village of Purulha. The locality is some 130 km east
of the closest populations of bromeliacia- group Chiropterotriton in Huehuetenango,
Guatemala, and is over 100 km west of the nearest known localities for C. nasalis

Review Committee for this Contribution
Robert L. Bezy
Arden H. Brame, Jr.

Richard Highton
John W. Wright

2Chesapeake Bay Center for Environmental Studies, Smithsonian Institution, Route 4, Box 622,
Edge water, Maryland 21037.

3Research Associate in Herpetology, Natural History Museum of Los Angeles County, 900 Ex-
position Boulevard, Los Angeles, California 90007, and Museum of Vertebrate Zoology, 2593
Life Sciences Building, University of California, Berkeley, California 94720.

2

Contributions in Science

No. 294

in Honduras. In 1972 the present authors had collected two species of Bolitoglossa
at the same locality where Uzzell found Chiropterotriton, but we did not encounter
salamanders of the latter genus on that visit, nor on a later trip (August, 1973).
Because Uzzell’ s specimens clearly represented an undescribed member of the genus,
we returned to the Purulha area a third time in November, 1974, in hopes of securing
additional material. On this occasion we collected a series of eleven Chiropterotriton .
We name this species for its provenance, a beautiful land of true peace:

Chiropterotriton veraepacis new species
Figure 1

Holotype: MVZ 1 12499. An adult male from 4.2 km (by road) S Purulha, Baja
Verapaz, Guatemala, collected by James F. Lynch, David B. Wake, and Theodore
J. Papenfuss, 14 November 1974. The holotype is one of a series taken at elevations
between 1740 and 1780 meters.

Paratypes: MVZ 112495-98 (4 specimens), LACM 123803 same data as holo-
type. MVZ 112490-112494 (5 specimens) collected at the type locality 13 November
1974. ANSP 28194-28197 (4 specimens) collected at the type locality in early 1973
by Thomas M. Uzzell, Jr.

Diagnosis: Compared to other Central American Chiropterotriton, C. veraepacis
is a rather small species, with an unusually long tail and narrow head. Eight sexually
mature individuals of both sexes measure 26.1-31.2 mm (mean 28.6 mm) SL4. The
tail is 1.26-1.43 times SL in undamaged adult males, whereas head width is 0.13-
0.14 times SL. Maxillary teeth are abundant (range 45-57; mean 49.0 for adults),
vomerine teeth are few (range 8-11; mean 9.6 for adults), and the nostrils are notably
enlarged (diameter equals 0.4-0. 6 mm in adults). C. veraepacis is distinguished from
C. nasalis by longer tail, much narrower head, smaller feet, and somewhat smaller
nostril5; fromC. picadoi by somewhat longer tail, narrower head, much longer limbs,
much larger feet, more maxillary teeth, and much larger nostrils; from C. bromeliacia
by somewhat smaller average size, narrower head, longer tail, and many more max-
illary teeth; from C. cuchumatanus by much longer tail, narrower head, more max-
illary teeth, and somewhat fewer vomerine teeth; from C. rabbi by smaller average
size, longer tail, much narrower head, and smaller nostrils; from C. xolocalcae by
much longer tail, much narrower head, more vomerine teeth, and much larger nostrils;
fromC. megarhinus by much longer tail, much narrower head, and somewhat smaller
nostrils. Other species of Chiropterotriton from Central America have a light inter-

4SL = standard length, here defined as the distance from the snout to the posterior angle of the
vent.

5 Statements concerning proportional distinctions between species are based on comparisons of
adult males at a common projected SL (30 mm). All character differences noted in the diagnoses
are significant at the 0.05 level or above.

1978

A new Guatemalan Chiropterotriton species

3

c.

Figure 1. Heads of adult
Chiropterotriton veraepacis,

males of species of Chiropterotriton, drawn to same scale, (a)
(b) Chiropterotriton bromeliacia, (c) Chiropterotriton picadoi.

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Contributions in Science

No. 294

orbital pigment band, at least in some specimens, but we have not seen any indication
of this feature in C. veraepacis . Both C. veraepacis and C. nasalis have nostrils that
are clearly visible by examination of the dorsal surface of the head from directly above
the specimen. In C. picadoi, C. richardi and members of the bromeliacia group the
nostrils are only slightly, or not at all, visible in dorsal view.

Description: To judge from the limited sample, C. veraepacis is a relatively small
member of the genus, the largest of fifteen specimens measuring 31.2 SL. The species
has the longest relative tail length and the narrowest relative head width of any of the
seven other northern Central American Chiropterotriton for which we have data (the
eighth known species from the region, the rarely collected C. barbouri of Honduras,
is not considered here). The number of maxillary teeth increases dramatically with
SL (r = 0.82; P< 0.01), but no size related increase in vomerine tooth count is evident
(r = 0.24; P > 0.1). The nostrils are moderately large in juveniles and subadults.
Nostrils increase in size in adults, contrary to what is seen inC. xolocalcae, C. rabbi,
and C. picadoi. The only adult female in the type series (MVZ 1 12497) has a shorter
tail and more maxillary teeth than have males of equivalent SL. Adult males possess
conspicuous oval- shaped mental hedonic glands.

Measurements of the holotype (in mm): Head width 3.9; snout to gular fold (head
length) 5.7; head depth at posterior angle of jaw 2.7; eyelid length 2.0; eyelid width
1.0; anterior rim of orbit to snout 1.8; horizontal orbit diameter 1.5; interorbital dis-
tance 2.3; distance between vomerine teeth and parasphenoid tooth patch 0.5; snout
to fore limb 7.9; distance separating internal nares 0.8; distance separating external
nares 0.8; snout projection beyond mandible 0.3; snout to posterior angle of vent (SL)
28.2; snout to anterior angle of vent 25.7; axilla to groin 14.3; tail length 39.8; tail
width at base 2.4; tail depth at base 2.6; fore limb length 6.7; hind limb length 7.6;
width of right hand 1.9; width of right foot 2.3.

Coloration (in life): The dorsum of the holotype is dark vinaceous brown. A red-
orange patch of pigment is present in the sacral region, but in contrast to the situation
seen in most Central American Chiropterotriton, there is no interorbital bar of light
pigment. A pair of obscure, red-brown patches of color are present in the otic region.
There is a hint of a jagged-edged dorsal stripe on the tail. The dense dorsal melanin
network is lighter laterally. A few small white iridophores are present at the side of
the head and trunk. The limb bases are light pink. The venter is slate gray, with the
tail somewhat darker than the belly and chin. Small white iridophores are scattered
on the chin, with fewer on the belly, and none on the underside of the tail. The animal
is fundamentally very dark, with other pigments (save at the tail base) becoming ev-
ident only on close examination.

The paratypes generally resemble the holotype in coloration. All fourteen para-
types are dark brown to black dorsally, all possess a light sacral patch (red, red-brown,
orange, or obscure red-orange), and all lack an interorbital bar of light pigment. Ven-
tral coloration shows more variation, ranging from medium gray to nearly black.
Contrast between the chin and the belly is greatest in those individuals with darker
bellies. The number of ventral white iridophores ranges from none to moderately
abundant, especially in the gular region. Most individuals show at least a few white

A new Guatemalan Chiropterotriton species

5

1978

flecks along the side of the head. In most individuals there is an obscure, irregularly
bordered caudal stripe, and in some it is faintly visible on the back as well.

Habitat: The type series was collected in an exceptionally humid cloud forest
which extends for several kilometers to the west and east of the type locality along
the north-facing slopes of the Sierra de Chuacus. According to the vegetational clas-
sification of Holdridge (1967), the type locality supports lower montane wet evergreen
forest (see also Savage, 1975). Large bromeliads, orchids, and other epiphytes are
conspicuously abundant, as are the tree ferns that are typical of cloud forests elsewhere
in Guatemala. Low palms are common in the understory of the forest. There are no
relevant weather data for nearby sites, but to judge from the aspect of the vegetation
and from the generalized precipitation map published by the Instituto Geografico
Nacional of Guatemala (1966), annual rainfall probably exceeds 3000 mm. The local
topography is extremely rugged, and much of the area is inaccessible to collecting
due to the extreme steepness of slope.

Ten of the eleven Chiropterotriton collected by the present authors were found
inside arboreal bromeliads ( Tillandsia and Catopsis spp) during daylight hours. The
eleventh specimen was encountered after dark on the upper surface of a palm leaf
approximately 1 m above the ground. Uzzell obtained the four ANSP paratypes and
an additional six specimens (ANSP 2885-90, too poorly preserved to measure) from
large bromeliads, 2-8 feet off the ground.

Other arboreal amphibians which occur at the type locality of C. veraepacis in-
clude two species of Bolitoglossa (a population tentatively assigned to B. helmrichi
and a less common, undescribed all-black species), Hyla bromeliacia, Plectrohyla
quecchi, and a large, fringe-limbed hylid similar to Hyla miliaria. All of the foregoing
species were found in arboreal bromeliads, but some occur under the loose bark of
downed logs as well. Terrestrial amphibians and reptiles collected in the vicinity of
the type locality include Eleutherodactylus brocchi, E. rugulosus, Anolis cobanensis,
A. crassulus, Barisia moreleti, Sceloporus taeniocnemis , and Thamnophis cyrtopsis.

The cloud forest inhabited by C. veraepacis became easily accessible in the sum-
mer of 1972 upon completion of an all-weather road through the mountains of Baja
Verapaz and Alta Verapaz. This previously undisturbed tract of forest is rapidly being
destroyed by lumbering and agricultural activities, and the prospects of continued
survival of C. veraepacis and other as yet undescribed endemic amphibians are bleak.

Range: Known only from the type locality and a nearby locality in Alta Verapaz,
Guatemala. To be sought in montane cloud forest along the humid, north-facing slopes
in the Chuacus-Minas mountain ranges of Baja Verapaz and Alta Verapaz. Thomas
Uzzell collected two specimens of Chiropterotriton in a bromeliad 30 feet above the
ground on the top of a ridge above Finca Volcan, Alta Verapaz (ANSP 28198-28199).
Stuart (1948) reports an altitude of 925 meters for the finca, and Uzzell’ s collecting
site is somewhat higher. Although these specimens are poorly preserved and cannot
be measured, we assign them to C. veraepacis on the basis of their general form,
large nostrils, light pigment spots at the base of the tail and absence of an interorbital
bar of pigment. C. veraepacis is the first species of salamander to be reported from
Baja Verapaz.

6

Contributions in Science

No. 294

Comparative Osteology

In our previous paper (Lynch and Wake 1975) we reviewed the osteological char-
acters of the five species that comprise the C. bromeliacia group. An additional five
species of Chiropterotriton occur south and east of the Isthmus of Tehuantepec: C.
veraepacis of Guatemala, C. nasalis and C. barbouri of Honduras, and C. picadoi
and C. richardi of Costa Rica (Wake and Lynch 1976). These ten species constitute
Chiropterotriton-beta.

In addition to the skeletal material for the bromeliacia group that was available
for our previous study, we have examined a second specimen of C. cuchumatamus .
A partial skeleton of the rare C. richardi provided limited information (Wake 1966),
and we have had one hand and one foot of an additional specimen (MVZ 99516). We
have examined one cleared and stained C. veraepacis, two C. nasalis, five C. picadoi,
and one specimen (ANSP 28200) of a population from the mountains south of Pueblo
Viejo, Depto. Yoro, Honduras, that is tentatively assigned to C. barbouri.

C. veraepacis differs sharply from all members of the C. bromeliacia group in
having prefrontal bones, in having frontal processes of the premaxillary bone which
arise separately and then fuse, separating again near their tips (Fig. 2), in having the

Figure 2. Three-quarter, anterolateral view of the nasal capsule region of three species of
Chiropterotriton, illustrating important features. Heavy lines, nasal bone; fine lines, intemasal
fontanelle; stipple, cartilaginous nasal capsule; blackened area, opening in nasal capsule for
external naris. Arrows point to area through which nasolacrimal duct passes ventrally into the
nasal capsule.

1978

A new Guatemalan Chiropterotriton species

7

fourth and fifth distal tarsals fused, and in having reduced numbers of carpal elements,
the result of ulnare-intermedium fusions (Fig. 3). In these characters it resembles both
C. nasalis and C. picadoi (Wake 1966, reported prefrontals to be present or absent
inC. abscondens = C. picadoi. This statement is based on one apparently pathological
specimen that has heavy ossification and in which the prefrontals and nasals are fused.
In the other four individuals examined the situation illustrated in figure 2 is seen.).
C. veraepacis, C. nasalis, andC. picadoi all lack septomaxillary bones and columellar
processes of the opercular apparatus, which are present in some of the bromeliacia
group. The new specimen of C. cuchumatamus has a pair of distinct septomaxillary
bones, so the elements have now been found in all members of the bromeliacia group
except C. xolocalcae.

In most osteological features C. nasalis and C. veraepacis are especially similar.
They differ from all members of the bromeliacia group in a number of derived char-
acters (the erratically structured, reduced nasal bones, the fused frontal processes of
the premaxillary, and the carpal and tarsal fusions). The members of the bromeliacia
group differ from these two species in one derived character, loss of the prefrontal
bones. Additional derived features of C. nasalis and C. veraepacis distinguish them
from some to most members of the bromeliacia group; these include loss of the sep-
tomaxillary bones and the preorbital processes of the vomers. C. nasalis has only a
rudimentary tibial spur, while the process is absent in the bromeliacia group. C. ver-
aepacis has a tibial spur.

The nasal capsules of C. nasalis and C. veraepacis have an orientation that dis-
tinguishes them from the members of the bromeliacia group and the other species
discussed here. The external nares are shifted so as to have a more dorsal orientation
than the nares of most species, which are typically directed anterolaterally in a nearly
vertical plane.

C. richardi andC. barbouri are the least well known species of Chiropterotriton -
beta, but we have some useful information. Both species have prefrontal bones,
preorbital processes of the vomers, and tibial spurs, but lack septomaxillaries and
columellar processes of the operculum. The ulnare and tibiale are fused in both, as
are distal tarsals four and five. Frontal processes of the premaxillary are fused for a
short distance above their origin in C. barbouri but not in C. richardi.

In the bromeliacia group the nasals are the main bony link between the frontals
and the maxillaries, but in C. nasalis, C. veraepacis and C. barbouri the prefrontals
provide that link. The nasals are reduced in the first two species. These features sug-
gest that enlarged nostrils have evolved in different ways in the two species assem-
blages, and that no constant set of characters accompanies nostril enlargement. Some
accommodation for the enlarged nostrils is required in the facial region of the skull,
but it may be the result, (a) of prefrontal loss and movement of nasals into the “pre-
frontal zone”, (b) of nasal reduction and increased prefrontal importance, (c) or of
both nasal and prefrontal reduction, but with both retaining important connecting func-
tions in the facial part of the skull (as occurs in C. richardi).

Both C. veraepacis and C. nasalis differ from C. picadoi and C. richardi in
lacking preorbital processes on the vomer and in having tear-drop shaped prefrontals
that are pierced for passage of the nasolacrimal ducts (Fig. 2). C. barbouri has preor-

Contributions in Science

No. 294

b.

Figure 3. Hands and feet of species of Chiropterotriton, drawn to same scale from cleared and
stained specimens with aid of microprojector. Cartilage stippled, (a) Left hand and (b) foot of
C. veraepacis; (c) foot of C. barbouri; (d) foot of C. nasalis; (e) hand, and (f) foot of C.
richardi; (g) foot of C. picadoi; (h) foot of C. bromeliacia.

1978

A new Guatemalan Chiropterotriton species

9

bital processes, but also has prefrontals very similar to those of C. veraepacis. The
nasal bones of C. veraepacis and C. nasalis are very reduced in size and irregular in
shape; the bones have weak or no articulations. Nasals are large, regular in form, and
well articulated in C. picadoi and C. barbouri. The small nostriled C. picadoi has a
very solid, strongly articulated skull, while the relatively large nostriled C. richardi
and the small nostriled C. barbouri have the more restricted articulation typical of
Guatemalan, Honduran, and Chiapan species of Chiropterotriton. In C. richardi the
nasals are rather small, but less reduced than in C. veraepacis and C. nasalis.

If the bromeliacia group is accepted as a monophyletic assemblage, the osteo-
logical data are consistent with a reasonably clear cladistic network for the eight spe-
cies we are able to consider in detail (Fig. 4). The characters used are as follows
(upper case letters indicate primitive states, lower case derived state; see Wake 1966,
for detailed analysis of characters):

A. Prefrontal and nasal form the bony connecting link between the frontal or the
center of the skull and the maxillary; the nasolacrimal duct passes between the
prefrontal and the nasal, a. No prefrontal bone; only the nasal bone is involved
in the link, and the nasolacrimal duct passes behind it. a'. Nasal bone reduced in
size and does not span frontal and maxillary; only prefrontal involved in link;
nasolacrimal duct pierces expanded ventrolateral part of prefrontal. States a and
a ' represent independent derivations from A in very different directions.

B. Septomaxillary present, b. Septomaxillary absent.

C. Frontal processes of premaxillary arise separately and do not fuse. c. Pro-
cesses fused at base but separate distally.

D. Distal tarsals 4 and 5 discrete, d. Distal tarsals 4 and 5 fused.

E. Vomer has a preorbital process, e. No process.

F. Intermedium and ulnare discrete in carpus, f. Intermedium and ulnare fused.

./ / /

$ aO

<r $

/ / / /

/ y y

H h

A, E

b,G B,g

a,e

A, b,c ,d,f ,g , H a , B,C , D, F,G , h

Chiropterotriton ft

Figure 4. Cladogram of eight species of Chiropterotriton- beta, constructed by use of osteo-
logical characters. The character states indicated apply to all species along the branch unless
modified states are indicated at a higher level in the dendrogram. See text for explanation.

10

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No. 294

G. Columellar process of operculum present, g. Process absent.

H. Tibial spur present, h. No tibial spur.

The first branching separates the bromeliacia group from all other species. The
second branching separates C. picadoi from C. veraepacis and C. nasalis on the basis
of loss of the preorbital process of the vomer and piercing of the prefrontals in the
latter two species. On osteological grounds C. nasalis and C. veraepacis are closest
to C. barbouri, and they are very distinct from the species of the bromeliacia group.
The osteological formulas for the poorly known species are, for C. barbouri: A (but
the nasolacrimal duct pierces the prefrontal as in a'), b, c (but the amount of fusion
is slight), d, E, f, g, H; for C. richardi: A, b, C, d, E, f, g, H. If placed on the
cladogram C. richardi would be on a branch derived from (but not a sister group of)
the main branch leading toC. picadoi, C. nasalis, andC. veraepacis. A branch lead-
ing to C. barbouri would be a more primitive, non-sister group of the C. vera-
epacis-C. nasalis line. In osteological formulae, barbouri is identical toC. picadoi,
but when one takes into account additional features, especially the shape of bones
such as the prefrontal and the route of the nasolacrimal duct, it seems clear that it is
more similar to its more osteologically derived neighbors than to C. picadoi.

The cladogram contains four convergences. We have rather arbitrarily coded C.
nasalis as h, even though a rudimentary tibial spur might be present. We have never
seen tibial spur rudiments in the bromeliacia group, but with larger samples it would
not be surprising to find an occasional rudiment. Septomaxillaries have not been ob-
served in C. xolocalcae, but experience has shown that these bones are found in at
least a small percentage of other members of the group, and the absence here may
reflect the small sample size. It is likely that the remaining two convergences (e and
g) are real.

In addition to the easily coded characters which have been used above, the gen-
eral arrangement and shape of the bones ofC. nasalis, C. veraepacis andC. barbouri
are similar, and in some details of foot structure (the shape of the terminal phalanges,
fig. 3) these species, C. picadoi and C. richardi are similar and differ from all mem-
bers of the bromeliacia group.

Morphometric Analysis
Methods

Lynch and Wake (1975) used a number of analytical approaches in comparing
the five species of the Chiropterotriton bromeliacia group. The number of specimens
of C. veraepacis, while small, is sufficient for some of the same kinds of quantitative
comparisons. Specifically, we have undertaken linear regression analysis of single
characters as they vary with SL, and multiple discriminant analysis of all characters
simultaneously in an attempt to clarify the phenetic and phyletic relationships among
eight of the ten Central American species of Chiropterotriton (C. bromeliacia, C.
cuchumatanus, C. megarhinus, C. nasalis, C. picadoi, C. rabbi, C. veraepacis, C.
xolocalcae ). C. barbouri, a poorly known species from western Honduras, and C.
richardi, a poorly known Costa Rican form, have been omitted due to lack of suf-
ficient comparative material. The reader is referred to our earlier paper for detailed

1978

A new Guatemalan Chiropterotriton species

11

description of statistical methodology and for discussion of the choice and scaling of
individual characters.

Analysis of individual characters : Eight characters are considered: standard
length (SL), head width (HW), nostril diameter (ND), combined length of hind limb
and fore limb (CL), foot width (FW), tail length (TL), number of maxillary teeth
(MT), and number of vomerine teeth (VT). As described in detail in Lynch and Wake
(1975), we have used linear regression techniques to project means and 95 percent
confidence limits for characters 2-8 to a common value of SL = 30 mm. To eliminate
the confusing effects of secondary sexual dimorphism and possibly nonlinear rates of
character change across very wide ranges of SL, only post-juvenile males (SL greater
than 24 mm) are included in the regression analysis.

Multivariate analysis of characters: The same eight morphological characters
examined in the regression analysis were utilized in the discriminant function analysis.
The purpose of this exercise was to define the combination of phenetic traits which
best distinguishes the Central American species of Chiropterotriton from one another,
and to eliminate the effects of intercorrelation among characters in assessing mor-
phological similarity among species. This latter aim is realized because discriminant
analysis defines statistically independent composite axes, each of which corresponds
to independently varying character complexes (see Blackith and Reyment 1971). The
Smithsonian Institution’s version of the SPSS program for stepwise discriminant anal-
ysis was used for all computations.

Phenetic clustering of taxa: The discriminant analysis defines the centroid of
each species cluster with respect to each of the seven discriminant axes which exist
for an analysis of eight groups and eight characters. Because the distances between
points in discriminant space (“D-space”) are not biased by intercorrelations among
characters, as is the case if phenetic distance is measured in simple Euclidian space,
it is often preferable to base phenetic clustering on the scores along the major dis-
criminant axes (for an example of the different results of these two metrics see Rob-
inson and Hoffmann 1975). The latter axes are said to define a “reduced space”
because most interspecific differentiation can be expressed using only a few of the
maximum possible number of discriminant axes. A two- or three-dimensional rep-
resentation of the group centroids (or of individual specimens) in discriminant space
often suffices as a summary of significant interspecific variation.

With the above in mind, we have used the generalized distance (=Mahalanobis
Distance) between the group centroids with respect to the first three discriminant axes
as the basis for phenetic clustering. The unweighted pair group method using arith-
metic averages (UPGMA), as described by Sokal and Sneath (1963) and Sneath and
Sokal (1973) was the clustering algorithm employed. The UPGMA technique is a
sequential, agglomerative, hierarchic, non-overlapping method (see Sneath and Sokal,
1973) which begins by defining a cluster (or several clusters) consisting of the least
dissimilar Operational Taxonomic Units (OTU’s), then adds individual OTU’s or other
clusters to pre-existing clusters on the basis of average phenetic similarity.

A second technique, the Prim algorithm (see Prim 1957; Farris 1970; Sneath and
Sokal 1973), was used to form a minimum-spanning network connecting the eight
OTU’s, again based on the generalized distance separating them. This algorithm min-

12

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No. 294

imizes the total length of the internodes connecting the OTU’s, and results in closest
phenetic neighbors always being linked by an internode. When superimposed on a
display of group centroids in reduced space, the Prim network provides a useful check
on the adequacy of the visual impression of inter-OTU distances (see Sneath and
Sokal, 1973: 255-256).

Results

Analysis of individual characters: Figure 5 and Table 1 summarize the results
of the regression analysis of single characters in post-juvenile males of the eight spe-
cies of Chiropterotriton. C. veraepacis has a longer tail and narrower head than the
other species in the comparison. The 95% confidence limits of the projected mean
values of TL and HW at SL = 30 mm show no overlap between C. veraepacis and
any of the remaining taxa. Some overlap is evident with respect to the other five
characters considered in the analysis, butC. veraepacis differs significantly (P<0.05)
from every other species in from three to six characters. The 95% confidence limits
for projected character values tend to be relatively broad for C. veraepacis and C.
nasalis, probably reflecting their small sample sizes.

A mosaic pattern of differentiation is found. For example, high maxillary tooth
counts may be associated with high vomerine tooth counts (C. xolocalcae ), moderate

species

tail length
30 36, 42 48

head width
40 ,4.5 , 49

combined
limb length
10, 12 14, 16,

foot width

,2.0 , , 30

number of
maxillary teeth
30 40 50,60

number of
vomerine teeth
10 14 18

nostril diameter
0.2 1.0 2.0

C. veraepacis

H-

■h

H-

4

-4.

4

■4

C. nasalis

4

4

"+■

■4

4

C. picadoi

■4

♦

4

4

4

4

C. bromehacio

4

4

4

4

4

4

4

C. cuchumatanus

4

+

4

H-

■4

4

C. megarhinus

4

4

4

4

•4

4

4

C. rabbi

4.

4

4

4

4

-4-

4

C. xolocalcae

*

4

4

4

4

4

4

1 1 1 III 1 1 1 l
30 36 42 48

in ri rr II i i

40 4.5 4.9

10 12 14 16

2 o' ' ' 'io

30 40 50 60

10 14 18

0 2 ' ' i o' '2 0

Figure 5. Predicted mean (vertical line) and 95% confidence limits (solid bar) for eight
characters in post-juvenile males of eight species of Chiropterotriton-beta. See text for further
explanation. Note that the column “head width” was incorrectly labeled “head length” in a
similar figure in Lynch and Wake (1975, fig. 7). All figures in mm.

Predicted mean character values and 95% confidence limits (i

1978

A new Guatemalan Chiropterotriton species

13

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14

Contributions in Science

No. 294

(C. nasalis ), or low (C. veraepacis) vomerine tooth counts. An exception to the pat-
tern of independent variation of characters is the invariable association of large nostril
size with low numbers of vomerine teeth (previously noted in the bromeliacia group,
Lynch and Wake 1975). C. picadoi is exceptional in having both few vomerine teeth
and very small nostrils. This species possesses a well-developed preorbital vomerine
process, as do the other small-nostriled species (C. rabbi, C. xolocalcae ) in the anal-
ysis, but the presence of the process is not always associated with a large complement
of vomerine teeth in other plethodontid salamanders (see Wake 1966; Lynch and Wake
1975: 34-35). Thus, our earlier conclusion that the preorbital vomerine process is
necessary, but not sufficient, for the presence of a large number of vomerine teeth,
is not affected by the pattern seen in C. picadoi.

Discriminant analysis: Figure 6 and Table 2 summarize the results of the stepwise
discriminant analysis of interspecific variation. The absolute magnitudes of the stan-
dardized discriminant coefficients (Table 2) are proportional to the relative importance
of each character in separating the species along each discriminant axis. The first three
discriminant axes, representing the first three discriminant functions, together account
for 95.7% of the observed interspecific morphological variation. Although the con-

Table 2

Summary of results of stepwise multiple discriminant analysis of variation
in post-juvenile male Chiropterotriton. Raw (unstandardized) and stan-
dardized coefficients are given for the first three discriminant functions,
which together account for 95.7% of the observed interspecific variation.
The magnitudes of the standardized coefficients are proportional to the
relative contribution of each variable on a given discriminant axis. The
most important variables on each axis are marked with an asterisk (*).

DISCRIMINANT FUNCTION
FIRST SECOND THIRD

CHARACTER

Raw

Stand.

Raw

Stand.

Raw

Stand.

SL

0.008

0.022

0.058

0.157*

-0.076

-0.204*

TL

0.003

0.012

0.029

0.128*

0.021

0.092

HW

-0.078

-0.042

-0.083

-0.045

-0.167

-0.090

CL

0.006

0.015

-0.147

-0.362*

-0.040

-0.100

FW

-0.076

-0.046

-0.060

-0.036

0.183

0.111

MT

0.001

0.011

-0.003

-0.028

0.036

0.350*

VT

-0.012

-0.060

-0.007

-0.038

-0.008

-0.041

ND

0.521

0.403*

-0.061

-0.047

-0.050

-0.038

CONSTANT

0.391

0.280

1.

,172

PER CENT
VARIATION

EXPLAINED 54.13 32.98 8.59

1978

A new Guatemalan Chiropterotriton species

15

Figure 6. Plot of the first three discriminant axes separating post-juvenile males of the species
of the Chiropterotriton bromeliacia group [bromeliacia (B), cuchumatanus (C), megarhinus
(M), rabbi (R), xolocalcae (X)] plus C. veraepacis (V), C. nasalis (N), and C. picadoi (P).
The dots are the positions of the group centroids for each species relative to these three variates.
The centroids are connected by a computed Prim network, which links the closest phenetic
neighbors. C. picadoi is about equally close to three species. See text for further explanation.

tributions of three of the remaining four discriminant axes are statistically significant
(P<0.05), their cumulative impact on the discriminatory process is small (4.3%). The
first discriminant function accounts for 54.1% of the interspecific variation; nostril
diameter (ND) is by far the most important determinant of the score along this axis,
as was also found to be the case in our earlier study of the bromeliacia group. The
second discriminant function (33.0% of interspecific variation) is influenced most
strongly by limb length (CL), and its most obvious effect is to sharply distinguish the
short-limbed C. picadoi from all other species. The third discriminant function (8.6%
of interspecific variation) shows high loading for maxillary tooth count (MT) and, to
a lesser extent, standard length (SL) and foot width (FW). This axis tends to separate
C. veraepacis and C. nasalis from C. picadoi and the bromeliacia group species.

16

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No. 294

Table 3

Generalized (Mahalanobis) distance values for centroids of eight OTU’s.
D = mean generalized distance.

SPECIES 2 3

1. C . bromeliacia 1.142 0.865

2. C. xolocalcae 0.402

3. C. rabbi

4. C. cuchumatanus

5. C. megarhinus

6. C. veraepacis

7. C. nasalis

8. C. picadoi

4

5

6

7

8

D

0.368

0.664

0.905

0.912

1.123

0.854

0.871

1.211

1.126

1.214

1.166

1.019

0.595

1.012

0.996

1.073

1.075

0.860

0.725

0.940

0.953

1.226

0.811

0.406

0.323

1.307

0.807

0.144

1.124

0.806

1.253

0.839

1.182

A three dimensional projection of the group centroids of each of the eight species
in relation to the first three discriminant axes (Figure 6) summarizes the differentiation
achieved by these axes. Viewed in D-space, C. picadoi appears to be the most isolated
of the eight species, mainly by virtue of its very high positive score along the second
discriminant axis. A close phenetic relationship between C. nasalis and C. veraepacis
is evident, and these two species appear more similar to C. megarhinus than to any
of the other bromeliacia group.

Table 3 gives the Mahalanobis distances separating the group centroids of the
eight species in the D-space defined by the first three discriminant axes. The impres-
sions gained from inspection of the three dimensional plot of group centroids are
strengthened by these distance estimates. The most similar pair of species is C. nasalis
and C. veraepacis (D = 0.14 units); no other species pair is separated by less than
0.32 units, the latter being the distance between C. nasalis and C. megarhinus. At
the other extreme, C. picadoi shows no strong similarity to any of the other species
(D = 1.18; range 1.07-1.31).

Despite the similarities of most of the species in gross appearance, the discrim-
inant analysis succeeded in assigning 95% of the 130 individual specimens to the
“correct” home population on the basis of morphometric criteria alone (Table 4).
This result confirms the utility of multivariate approaches in making morphological
comparisons among OTU’s which show complex patterns of overlap in single char-
acters and which exhibit size-related variation in most morphological traits. Of the
seven misidentifications which occurred, three were assignments of C. bromeliacia
and C. rabbi to C. cuchumatanus, three were assignments of C. rabbi to C. xolo-
calcae, or vice versa, and one is a misclassification of C. nasalis as C. megarhinus .
In spite of the small overall phenetic separation of C. nasalis and C. veraepacis in-
dividuals of both species were not mismatched by the discriminant procedure.

Phenetic clustering; The Prim network linking the eight Chiropterotriton species
is superimposed over the three-dimensional projection of the group centroids in figure
6. The distortion resulting from the geometry of the projection and from the omission

1978

A new Guatemalan Chiropterotriton species

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Table 4

Classification matrix based on discriminant analysis of post-juvenile male
Chiropterotriton from eight populations. Entries along the main diagonal
are “correct” matches of individuals to their source population based on
eight morphological characters. See text for further explanation.

PREDICTED GROUP MEMBERSHIP

1

2

3

4

5

6

7

8

Actual Group

N

1. C. bromeliacia

33

32

0

0

1

0

0

0

0

2. C. xo locale ae

22

0

21

1

0

0

0

0

0

3. C. rabbi

22

0

2

18

2

0

0

0

0

4. C. cuchumatanus

9

0

0

0

9

0

0

0

0

5. C. megarhinus

13

0

0

0

0

13

0

0

0

6. C. veraepacis

8

0

0

0

0

0

8

0

0

7. C. nasalis

7

0

0

0

1

0

0

6

0

8. C. picadoi

16

0

0

0

0

0

0

0

16

of the fourth and higher discriminant axes evidently is small, for the Prim linkage of
OTU’s which show the highest phenetic similarity conforms well with the visual
impression of similarities gained from the projection. The network is basically a chain
which connects seven of the species in the order xolocalcae-rabbi-cuchumatanus-
bromeliacia-megarhinus-nasalis-veraepacis . The eighth species, C. picadoi, lies well
off this main sequence, and is about equally as distant from C. rabbi, C. bromeliacia,
and C. nasalis.

The UPGMA phenogram, based on the same matrix of generalized inter-OTU
distances used to construct the Prim network, is shown in figure 7. Summarizing, we
see a tight cluster formed by C. veraepacis and C. nasalis, which is connected to
the‘ ‘looser” grouping of four bromeliacia group species by C. megarhinus, the fifth
species in that group. C. picadoi shows slightly more similarity to the small-nostriled
members of the bromeliacia group than to the other OTU’s, but it is not very similar
to any of the species. The linkage pattern among the five species of the bromeliacia
group produced by the UPGMA analysis is identical to that derived previously (Lynch
and Wake 1975: Fig. 10a) from an 11 -character phylogenetic clustering algorithm,
the WISS method of Farris, Kluge, and Eckhardt (1970).

DISCUSSION

The species of Chiropterotriton considered in this paper have several ecologically
important features in common. All are small, arboreal or crevice-dwelling salaman-
ders which inhabit wet montane forest, and all have very limited geographic ranges.
Sympatry between congeners is unknown in northern Central American Chiroptero-

18

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No. 294

Figure 7. UPGMA phenogram of eight species of Chiropterotriton based on Mahalanobis
Distance between group centroids with respect to the first three discriminant axes. (B) C.
bromeliacia; (C) C. cuchumatanus; (M) C. megarhinus; (N) C. nasalis; (P) C . picadoi; (R)
C. rabbi; (V) C. veraepacis; (X) C. xolocalcae. See text for futher explanation.

triton, although co-occurrence of several pairs of species of Chiropterotriton- alpha
has been documented north of the Isthmus of Tehuantepec in the Sierra Madre Ori-
ental. In addition, C. picadoi and C. richardi of the beta assemblage are sympatric
in Costa Rica. In the absence of a sympatry test, one can always argue that the isolated
populations of Chiropterotriton in Central America do not merit full specific rank.
However, the unique combination of external morphometric traits which characterizes
each of the Central American populations, combined with the osteological differences
between most of the forms, makes it highly likely that these populations do in fact
represent separate evolutionary units. We are actively investigating genetic divergence
between species of tropical salamanders, and results to date lend support to our con-
clusions based on morphology.

The data from morphometries and from osteology produce somewhat conflicting

1978

A new Guatemalan Chiropterotriton species

19

pictures of evolutionary relationships. The UPGMA clustering technique based on the
external morphology and dentition places C. nasalis and C. veraepacis well within
the bromeliacia group. These seven species share more in common ecologically than
they do osteologically. All are small species that occur in basically arboreal sites in
cloud forests. They appear to be ecomorphs, with similar, adaptive structural re-
sponses to similar selective pressures. Certainly one reasonable hypothesis is that they
are close relatives, derived from a common stock that was, itself, adapted to climbing
in cloud forest habitats.

On osteological grounds this suggestion does not appear to be valid. An alternate
hypothesis postulates an early separation between the ancestors of the bromeliacia
group and the ancestors of all other Chiropterotriton- beta. These two lineages are
sister groups, in the sense of Hennig (1966), with both lineages displaying both prim-
itive and derived characters. The bromeliacia group has remained in Nuclear Central
America, and has speciated but not undergone much additional differentiation. In con-
trast, the other assemblage is rather diverse, and widespread. C. veraepacis and C.
nasalis are close relatives, distinct from each other in morphometric features but not
in osteology. The cladistic species pattern of figure 4 leads us to suggest that mor-
phological similarity of these two species to the bromeliacia group results from con-
vergence by different stocks in response to similar selective pressures.

On osteological grounds both C. richardi and C. barbouri seem allied with C.
picadoi, C. nasalis, and C. veraepacis rather than with the bromeliacia group. C.
barbouri is a relative of C. nasalis and C. veraepacis and is somewhat intermediate
between these two species and C. picadoi in osteological structure. Possibly C. ri-
chardi is the earliest known derivative of a Chiropterotriton -beta stock. It is the only
species in Central America with any degree of resemblance to members of the genus
Oedipina, lives sympatrically with species of that genus, and, like Oedipina, has been
found living in holes in moss-covered earth banks. This elongate, diminutive animal
with very short limbs and a long tail, closely resembles juvenile Oedipina in habitus.
Possibly it is a semi-fossorial form.

A general character state formula for the genus Oedipina is: a, b, c. d, E, f, g,
h (one population of one species of Oedipina appears to have septomaxillaries but b
rather than B is used because of the clear preponderance of that state). This formula
is compatible with the hypothesis that an animal rather like C. richardi might have
given rise to Oedipina . Further suggestion of relationship comes from details of the
structure of the feet of C. richardi and Oedipina. In addition to having the carpal and
tarsal fusions typical of the nasalis group (character states d, f), C. richardi also has
a fusion of the centrale and distal carpal 4, (Fig. 3). Oedipina also typically has all
of these fusions, and fusion of distal carpal 4 and the centrale has also been reported
in Parvimolge toxvnsendi and various species of Thorius (Wake 1966). Fusion of the
tibiale and centrale occurs in the tarsus of C. richardi, but has not been encountered
elsewhere.

The relationship of C. picadoi to other species of Chiropterotriton -beta is ob-
scure. On the basis of sharing many derived states with C. barbouri, C. nasalis, and
C. veraepacis (b, c, d, f, g) we are tempted to postulate close relationship. Yet, C.

20

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No. 294

picadoi is very different from these three species in external morphological features
and dentition. It has very short limbs and small feet, and in many features resembles
the even more elongate and short-limbed C. richardi.

We earlier (Wake and Lynch 1976) assigned C. nasalis, C. barbouri, C. ri-
chardi, and C. picadoi to a nasalis group, thus placing all Chiropterotriton- beta in
but two groups. We retain that arrangement here, but note that the nasalis group is
morphologically far more diverse than the bromeliacia group.

Zoogeographic patterns conform well with our hypothesis concerning evolution-
ary relationships. The bromeliacia group is a cluster of species with high phenetic
similarity sharing many derived osteological characters. They are so far known from
a number of localitites in western Guatemala and Chiapas. The nasalis group is far
more diverse than the bromeliacia group in morphometric and osteological traits.
When additional information becomes available it may be possible to subdivide the
nasalis group into three parts: 1) C. nasalis, C. barbouri, andC. veraepacis, a mainly
northeastern group of arboreal species from Guatemala and Honduras that seems well
defined osteologically; 2) C. picadoi of Costa Rica, allied to the former three species
by derived osteological traits but not by ecology or external morphology; 3) C. ri-
chardi of Costa Rica, a highly distinctive, diminutive, elongate species with a com-
bination of generalized (skull) and specialized (feet) osteological traits that distinguish
it from all other species of Chiropterotriton .

The recent (February, 1976) Guatemalan earthquake has led to renewed interest
in the structure and history of Nuclear Central America, one of the most complex and
seismically active regions of the world. As a result of detailed investigations of land
movements during the earthquake, the principal faults have been sharply defined, and
the history of the region has been placed in new perspective. The Cocos plate is
moving from the southwest and being subducted where it meets the Caribbean and
North American plates in what Plafker (1976) calls the Middle American Megathrust.
The North American plate is moving mainly westward, and the Caribbean plate is
being forced eastward by the combined plate movements. The Motagua fault, clearly
outlined by the recent earthquake, lies at the border of the Caribbean and North
American plates. The extreme western end of the Caribbean plate is being “squeezed”
between the Cocos and North American plates, and Plafker speaks of this region as
being “decoupled” from the plate. This is the area of maximum volcanic activity in
southern Guatemala. The amount of movement along the Motagua fault has been
great, with a probable minimum movement of 200 km since Miocene being estimated
by Plafker. Estimates of total movement along the fault range from a few hundred to
over 1000 km. The movements are great enough and recent enough to have had pro-
found impact on salamander distribution and evolution in the area. Chiropterotriton,
for example, is restricted to montane and lower montane cloud forests in Central
America. Low mountain ridges of nearly continous cloud forest are required for dis-
persal of these salamanders. In the present instance, we have shown thatC. veraepacis
and C. nasalis are similar in structure and we think that they are close relatives. C.
veraepacis is found in a region that lies a little over 100 km west of the nearest
population of C. nasalis. However, C. veraepacis occurs to the north of the Motagua
fault, right along a branch of the Polochic fault, on the edge of the North American

1978

A new Guatemalan Chiropterotriton species

21

plate, while C. nasalis occurs to the south and east of the fault zone, on the Caribbean
plate. Thus there is no continuous montane link between the two areas now, and it
is extremely unlikely that one has extended across the fault in the past. It is more
likely that the two species populations have dispersed to their present sites by routes
extending from the west in both instances. Presumably these populations have been
separated for long periods of time.

Chiropterotriton- beta is sufficiently different from Chiropterotriton- alpha that
the erection of a new genus for the former group might be justified. We choose not
to do so at this time, for Chiropterotriton -beta is so diverse in structure that it would
be difficult to diagnose meaningfully. Further, the osteo logical evidence presented
here suggests that even a finer division of Chiropterotriton may be required than a
simple two-way split. The key species in any future taxonomic revision of this group
is C. richardi, a highly specialized species that might be derived from an early stock
that gave rise not only to the remainder of Chiropterotriton -beta but also Oedipina.

The present study supports the view that a great amount of evolutionary differ-
entiation has occurred in the lineages of plethodontid salamanders in the New World
tropics.

ACKNOWLEDGMENTS

This research has been supported by the National Science Foundation (current
grant BMS 74-20922, D. B. Wake, Principal Investigator). Thomas Uzzell gener-
ously provided information and lent us important specimens, for which we are most
grateful. We continue to benefit from the counsel of L. C. Stuart in our work on
tropical salamanders. Figure 1 was executed by Samuel S. Sweet, and Gene M.
Christman aided in the preparation of the other illustrations. Pedro Alberch prepared
the Spanish summary.

RESUMEN

En este trabajo se describe una diminuta nueva especie de salamandra pleth-
odontida de grandes narinas, Chiropterotriton veraepacis, que habita en bromeliaceas
en la zona de bosque nublado montano al sur de Puruhla, Baja Verapaz, Guatemala
y en las zonas cercanas de Alta Verapaz, Guatemala. C. veraepacis muestra seme-
janzas en muchos caracteres feneticos con C. nasalis del nordeste de Honduras, que
es una especie de cola mas corta, cabeza mas ancha y narinas de mayor tamano. La
nueva especie se asemeja en ecologfa y algunos caracteres estructurales a las especies
del grupo bromeliacea de Guatemala y Chiapas, pero difiere de ellas en caracteres
osteologicos.

A pesar de que C. nasalis y C. veraepacis son muy distintos en la mayoria de
caracteres morfologicos y ecologicos de la especie costarricense C. picadoi, estas
especies son parecidas en osteologia. Tambien se presenta informacion adicional re-
ferente a las poco conocidas especies, C. richardi (Costa Rica) y C. barbouri (Hon-
duras). Asimismo se tratan las posibles relaciones entre las especies de este genero.

22

Contributions in Science

No. 294

LITERATURE CITED

Blackith, R. E. and R. A. Reyment. 1971. Multivariate Morphometries. Academic Press,
London and N.Y., ix + 412 pp.

Farris, J. S. 1970. Methods for computing Wagner Trees. Syst. Zool. 16: 83-92.

Farris, J. S., A. G. Kluge and M. J. Eckhardt. 1970. A numerical approach to phylogenetic
systematics. Syst. Zool. 19: 172-189.

Hennig, W. 1966. Phylogenetic Systematics. Trans. D. D. Davis, R. Zangerl. Univ. Illinois
Press, Urbana. 263 pp.

Holdridge, L. R. 1967. Life zone ecology (2nd ed.) Tropical Sci. Center, Costa Rica. 206 pp.

Instituto Geografico Nacional (Guatemala). 1966. Atlas preliminar de Guatemala.

Lynch, James F. and D. B. Wake. 1975. Systematics of the Chiropterotriton bromeliacia
group (Amphibia: Caudata), with description of two new species from Guatemala. Los
Angeles County Nat. Hist. Mus. Contrib. Sci. 265: 1-45.

Plafker, G. 1976. Tectonic aspects of the Guatemala earthquake of 4 February 1976. Science,
193: 1201-1208.

Prim, R. C. 1957. Shortest connection networks and some generalizations. Bell System Tech.
J. 36: 1389-1401.

Robinson, J. W. and R. S. Hoffmann. 1975. Geographical and interspecific cranial variation
in big-eared ground squirrels ( Spermophilus ): a multivariate study. Syst. Zool. 24: 79-188.

Savage, J. M. 1975. Systematics and distribution of the Mexican and Central American stream
frogs related to Eleutherodactylus rugulosus. Copeia, 1975: 254-306.

Sneath, P. H. A. and R. R. Sokal. 1973. Numerical Taxonomy. Freeman and Co., San
Francisco: xv -I- 573 pp.

Sokal, R. R. and P. H. A. Sneath, 1963. Principles of Numerical Taxonomy. Freeman and
Co., San Francisco, xvi -I- 359 pp.

Stuart, L. C. 1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. Publ.
Mus. Zool. Univ. Michigan, 69: 1-109.

Wake, D. B. 1966. Comparative osteology and evolution of the lungless salamanders, family
Plethodontidae. So. Calif. Acad. Sci., Mem. 4: 1-111.

Wake, D. B. and J. F. Lynch. 1976. The distribution, ecology, and evolutionary history of
plethodontid salamanders in tropical America. Bull. Nat. Hist. Mus. Los Angeles Co. 25:
1-65.

Accepted for publication October 27, 1976.

SOI- 73

cau^s

NUMBER 295
MAY 15, 1978

FISHES OF THE SANTA CLARA RIVER SYSTEM,
SOUTHERN CALIFORNIA

ft

0A HSOW/^jT

MAY 2 2 1978

By Michael A. Bell

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTDNS IN SCENC6

Published by the NATURAL HISTORY MUSEUM
OF LOS ANGELES COUNTY
900 Exposition Boulevard, Los Angeles, California 90007

SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

Prior to November 30, 1973, publications of the Natural History Museum have appeared
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CONTRIBUTIONS IN SCIENCE contain articles in the earth and life sciences, presenting
results of original research. Emphasis is intended principally for papers allied to biosystematic
research, but other subjects and review-oriented ones will be considered. Number 1 was issued
on January 23, 1957. Contributions must be not less than 8 nor exceed 72 printed pages.

INSTRUCTIONS FOR AUTHORS

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MANUSCRIPT FORM.— (1) In preparation of copy follow the 1972 CBE Style Manual,
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should be sent to the Editor, Museum Publications, Natural History Museum of Los Angeles
County, 900 Exposition Boulevard, Los Angeles, California 90007.

Printed in the United States of America by Chapman's Phototypesetting on 70# Patina Book

FISHES OF THE SANTA CLARA RIVER SYSTEM,
SOUTHERN CALIFORNIA1

By Michael A. Bell2

Abstract: The distribution of fishes within the Santa Clara River and most of
its tributaries in which fishes were found is described from field observations and 46
fish collections. Fifteen species of fishes were collected, of which five probably are
native to the system. All native species are peripheral freshwater fishes so there is
no evidence of faunal exchange with adjacent river systems. Gasterosteus aculeatus
and Gila orcutti occurred at almost all collecting stations while five species occurred
at only one. This river system contains a high diversity of freshwater fishes for south-
ern California streams. The Santa Clara River system is vulnerable to habitat destruc-
tion by urbanization, and potential threats to fishes of the system are discussed.

INTRODUCTION

Although many papers have discussed individual fish species from the Santa
Clara River system (Girard 1854; Hubbs 1967; Hubbs, Hubbs and Johnson 1943;
Miller 1960, 1968, 1972, 1973; Ross 1973; Smith 1966), its fish fauna has not been
described previously. By virtue of its proximity to Los Angeles, this drainage currently
is subject to rapid urbanization with the attendant dangers of habitat destruction and
pollution. The only known native population of the endangered unarmored threespine
stickleback, Gasterosteus aculeatus williamsoni (Girard), still survives in this drain-
age. This drainage also harbors some introduced fishes endemic to southern California
and some species of game fishes. Thus, it is desirable to describe the fish fauna before
disturbance of the ecosystem causes the elimination of some species. This description
will serve in the future as a base line to assess the impact of urbanization on the fish
fauna.

MATERIALS AND METHODS

Forty-six fish collections were made at varying intervals from the mouth of the
Santa Clara River system to as far upstream as water existed except in Piru and Santa
Paula creeks primarily between September 4 and November 9, 1975 (Fig. 1). Col-
lections were made in the Santa Clara River, Todd Barranca, Sespe Creek, Piru Creek

!Review Committee for this Contribution
Robert J. Lavenberg
J. D. McPhail
Robert R. Miller
Camm C. Swift

2Research Associate in Ichthyology, Natural History Museum of Los Angeles County, and Depart-
ment of Biology, University of California, Los Angeles, California 90024.

2

Contributions in Science

No. 295

i

Figure 1. Map of the Santa Clara River system- showing the location of collections (numbers) and observations (letters).

1978

Fishes of Santa Clara River

3

and San Francisquito Canyon. Arrastre Canyon and Santa Paula Creek were examined
briefly, but no collections were made. The original purpose of the sampling program
was to determine the distribution of threespine stickleback {Gasterosteus aculeatus )
phenotypes. This species favors areas of low velocity flow so most collections were
made in backwaters, side streams, pools and mainstream margins. Samples are thus
biased against those species preferring rapidly flowing water. For this reason, esti-
mates of abundance, which may be misleading were not made, and the absence of
a species from a sample should be interpreted with caution.

Collecting stations were selected for accessability and the presence of suitable
habitat for Gasterosteus . The position of stations was determined in the field using
local landmarks, and they were recorded on U.S.G.S. 7.5 minute series (topographic)
maps. The distance of collection stations from the mouth in the Santa Clara River and
from the confluence with the Santa Clara River in tributaries was determined by step-
ping off that distance on the topographic maps using dividers set at 0. 1 km. Stream
gradient was determined by stepping off the distance between one or two contour lines
up and downstream of the station with dividers set for 500 or 200 feet. Because of
irregularities in stream course, the distance from the mouth or confluence tends to be
a slight underestimate and the gradient a slight overestimate using this method.

Surface temperature and velocity were measured where the majority of fishes
was collected at each station. Thermometers were calibrated to within 0.5 C of the
freezing and boiling point of distilled water. Water temperature tended to vary with
the air temperature. Water velocity was determined by measuring the time required
for a vial partially filled with water (so only a comer protruded above the surface) to
drift 5 m. Water depth, stream width, amount of vegetation, bottom composition and
water color were typical of the habitat, but collecting frequently extended across a
heterogeneous segment of stream.

Fishes were collected with a 10 foot (3.048 m), Vs inch (3. 175 mm) mesh knitted
nylon seine. The collecting effort usually varied inversely with the abundance of Gas-
terosteus. Specimens were fixed in 10% formalin and transferred to 50% isopropyl
alcohol 4 to 10 days after collection. Eddy (1957), Kimsey and Fisk (1960), Robins
and Miller (1957) and Smith (1966) were useful for identification. The current sci-
entific and common names were verified in Baily (1970). All collections were de-
posited in the fish collection of the Natural History Museum of Los Angeles County
(LACM). These are station 1, LACM 34071, station 2 to 44, LACM 34198 to 34240
respectively, station 45, LACM 35228, and station 46, LACM 34241. LACM 35227
also came from station 31 and LACM 35648 was collected near station 30. Additional
specimens from the California Academy of Sciences Fish Collection (CAS) were ex-
amined to verify early collection records (CAS 20283, CAS 20284 and CAS Acc.
No. X:30).

I attempted to determine whether species presently occuring in or previously re-
ported from the Santa Clara River system were native or introduced. This is a rela-
tively simple task for species known to be introduced from distant sources, but those
native to southern California present a problem. One must depend on historical rec-
ords, distribution patterns, the original presence of appropriate habitats and the exis-
tence of dispersal routes for entry into the system. Only negative historical data can

4

Contributions in Science

No. 295

be brought forth to support the hypothesis that a species is introduced. The failure of
a species to be observed or collected (negative data) until recent times may be a result
of recent introduction or from erroneous observation, identification or incomplete col-
lection in the past. Even if there is a record of introduction, the species may already
have been present before the introduction. The confidence placed in the conclusion
based on historical data that a species is introduced depends on the quality of the
original observation. Reasoning based on distribution patterns, the presence of ap-
propriate habitats and dispersal routes is inferential. Fishes do not always occur in
habitats that they can disperse into and that appear to be appropriate for them. Thus,
as much evidence as possible must be brought to bear on the question of whether a
fish species is native, and the answer obtained may never be satisfactory.

RESULTS

Description of Streams

The Santa Clara River system is composed of the Santa Clara River and a large
number of tributaries primarily draining from the north (Fig. 1). The drainage is
bounded on the southwest by the Santa Susana Mountains and on the southeast by the
San Gabriel Mountains. No substantial tributaries of the Santa Clara River drain these
mountain ranges. The Santa Susana Mountains are drained to the south by Calleguas
Creek and its tributaries. The southern slope of the San Gabriel Mountains is drained
by tributaries of the Los Angeles and San Gabriel Rivers.

In the west, the headwaters of Sespe and Piru creeks are interdigitated with those
of the Cuyama River, a tributary of the Santa Marie River and tributaries of the small
Ventura River system.

Drainage north of the Santa Clara River system is by creeks which eventually
disappear into the southern San Joaquin Valley. The eastern comer of the Santa Clara
River system is bounded by these creeks as well as those of the Los Angeles River
system and some draining into the Mojave desert.

The Santa Clara River, Santa Paula, Sespe, Piru and Castaic creeks, San Fran-
cisquito and Arrastre canyons and Todd Barranca were studied. Other tributaries east
of Saugus were not studied because J. N. Baskin (personal communication) reported
no fishes there. Some tributaries in the western portion of the drainage could not be
studied or received cursory examination because of limitations of time, funds or
access.

The Santa Clara River was examined from its headwaters to its mouth. Flow is
intermittent over substantial lengths of the stream (dotted lines, Fig. 1) and the geo-
graphic extent and duration of desiccation varies yearly. The section between Lang
and Saugus is dry except during heavy downpours, a condition that apparently has
existed since at least the middle of the last century (Miller 1960). The Santa Clara
River and its tributaries are subject to flooding some winters. The river flows through
a broad, primarily sandy -bottomed valley. In most places, the flood plain is lined by
earth and rock or wire and debris barriers. The Southern Pacific Railroad runs the
length of the Santa Clara River, built across the flood plain on elevated grades in
places. Upstream of Saugus the flood plain is occupied by gravel pits, small recrea-

1978

Fishes of Santa Clara River

5

tional parks or is undeveloped. The Saugus area presently is undergoing rapid urban-
ization. The flood plain, beginning just upstream of Piru Creek, is occupied by citrus
orchards, which bound most of the river on both sides except for small gaps at the
cities of Fillmore, Santa Paula, Satacoy and Oxnard. The Oxnard-Satacoy region also
is an area of rapid urban growth.

The first tributary studied was Todd Barranca, a small stream that drains Wheeler
Canyon and flows across the flood plain of the Santa Clara River, entering it at Santa
Paula. Fishes were present in the lower portion.

Only the lower kilometer of Santa Paula Creek, which is enclosed in a rock and
earth levee was examined (stat. C). No fishes were seen.

Sespe Creek is a large tributary of the Santa Clara River. Numerous small wa-
terfalls a few centimeters to more than 3 m high occur from 12 to 17 km (and probably
farther) upstream of the confluence with the Santa Clara River (stats. 32-35). Much
of the stream flows through deep, narrow canyons over rocky substratum. Practically
the whole Sespe Creek drainage is contained in Los Padres National Forest (including
the Sespe Condor Sanctuary, where no samples could be collected) and is protected
as recreational land or wildlife sanctuary.

Piru Creek is the largest tributary to the Santa Clara River. Flow volume fluc-
tuates according to the amount of water released from Lake Piru through Santa Felicia
Dam, and the creek receives imported water from Pyramid Reservoir. The canyon
through which Piru creek flows in that portion studied is broad and sandy-bottomed.
Piru Creek is primarily within the Los Padres National Forest.

Castaic Creek is located in a dry, broad, sandy-bottomed valley. Castaic Lake
is a reservoir that receives imported water from Pyramid Reservoir. The only flowing
water seen in this creek during the summer was water being released from a small
impoundment below Castaic Lake at station 45.

San Francisquito Canyon contains three areas of continuous flow: where it joins
the Santa Clara River, and at two points upstream. One point upstream is where water
released from Drinkwater Reservoir tumbles out of Drinkwater Canyon and flows for
about 200 m along the bottom of San Francisquito Canyon before sinking into the
sand. On February 1, 1976 this stream segment had increased to 1400 m long. Such
extensions are characteristic of the Santa Clara River system in the winter. The second
point upstream where the stream flows is for a few kilometers below San Francisquito
Powerhouse No. 1 . The canyon bottom varies in width and generally is dry and sandy.

Arrastre Canyon is one of several canyons that converge to form the headwaters
of the Santa Clara River. A small flow descends the steep sandy bottom of this shallow
canyon, sinking into the sand and forming small pools. Like San Francisquito Canyon,
this is a remote and little disturbed canyon.

Fish Distributions

Fifteen fish species were collected from 46 collecting stations. The characteristics
of these stations are indicated in Table 1 . Fishes collected are listed below (numbers
in parentheses indicate the number of stations at which the species was collected):

Gasterosteus aculeatus Linnaeus Threespine stickleback (42)

Gila orcutti (Eigenmann and Eigenmann) Arroyo Chub (37)

6

Contributions in Science

No. 295

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Gambusia ajfinis (Baird and Girard) Mosquito fish (24)

Catostomus (. Pantosteus ) santaanae (Snyder) Santa Ana sucker (20)

Pimephales promelas Rafinesque Fathead minnow (15)

Salmo gairdneri Richardson Rainbow trout (5)

Lepomis cyanellus Rafinesque Green sunfish (6)

Micropterus salmoides (Lacepede) Largemouth bass (4)

Ictalurus punctatus (Rafinesque) Channel Catfish (1)

Cottus asper Richardson Prickly sculpin (3)

Notepiigonus crysoleucas (Mitchill) Golden shiner (1)

Dorosoma petenense (Gunther) Threadfin shad (1)

Eucyclogobius newberryi (Girard) Tidewater goby (1)

Leptocottus armatus Girard Pacific staghorn sculpin (1)

Lampertra tridenteta (Gairdner) Pacific lamprey (2)

Fish collection records are summarized in Table 2 and the locations of collection
stations are shown in figure 1 .

Gasterosteus acculeatus occurred in every stream where fishes were found except
in Arrastre Canyon and Castaic Creek. It was abundant at all stations where it occurred
except in the Santa Clara River between Saugus and Piru and in the headwaters of
some tributaries. This species was collected by J. N. Baskin (personal communication)
at my station F, farther upstream in San Francisquito Canyon than I was able to find
it.

Gila orcutti penetrates farther into headwaters than Gasterosteus . In addition to
being widespread in the Santa Clara River and all tributaries in which fishes were
found, it was observed in Arrastre Canyon (stat. G) and farther up San Francisquito
Canyon (stat. F), Sespe Creek (stat. D) and Piru Creek (stats. 43, 44, E) than
Gasterosteus .

Gambusia ajfinis, Catostomus santaanae and Pimephales promelas are all wide-
spread in the Santa Clara River but are restricted in some tributaries. Gambusia ajfinis
apparently is absent above station 22 in the upper Santa Clara River and was not found
above station 30 in Sespe Creek or station 39 in Piru Creek. Generally the absence
of Gambusia is associated with an increase in stream gradient (Table 1). Catostomus
santaanae was absent from small tributaries like Todd Barranca, San Francisquito
Canyon and Arrastre Canyon and also from the headwaters of Sespe Creek (above
stat. 32). However, J. N. Baskin (personal communication) found it and Gila orcutti
in isolated pools in Mill Canyon, which joins the Santa Clara River near station 23.
Pimephales promelas has a more restricted distribution. In the Santa Clara River, it
occurred below station 14 and has only entered the lower gradient portions of Sespe
Creek, below station 30. However, fathead minnows were seen above Lake Piru and
were collected in the lake, in two stations downstream in Piru Creek and below Castaic
Lake.

The only other widespread species is Lepomis cyanellus, which was taken at six
disjunct stations in the Santa Clara River, Todd Barranca, Castaic Creek and seen in
Sespe Creek. It probably occurs elsewhere, but may be rare and difficult to collect.

Other species found in the system are either locally abundant or associated with
unusual conditions. Salmo gairdneri lives in the discharge of Fillmore Fish Hatchery

Table 2

Distribution of fishes in the Santa Clara River systems. Symbols: +, species collected; S, species seen, but not collected.

1978

Fishes of Santa Clara River

9

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sinqoSojjfong

asuaudidd

VWOSOAOQ

svjndjOsLi.i

snuoSiiudjOjq

Addsv

snuoj

smviJund

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^ sdpiouqvs

[ ^ snAdidoAJij/\j

MJ

kj

a, snijduunxj

siwoda 7

!AdUpAU?8

ouqvs

svjduiOAd

sdjvqddiuic]

dDunmuvs

smuoisojDj

smgv

vjsnquivQ

UIYIDAQ

VUD

+

+

+

• • • + +

+ • + + + +

•••• + •

+ + + + + +

+ + • + + +

+

. ... GO •

+ • + • • +

+ • + + + +

+ • • + • +

+ + + + 00 +

+

• + •+ + +

+ + + + + +

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smud/ruv

snaisoAdisnp)

+ + + + + + +

+ + + +

+ + +

W 1 H
Oi =3 <
H H
oo GO

I 00

(Nr0'Nj-i/~)\0t-~000''O — (N (O >0 'O I — OO

Table 2 (continued)

Distribution of fishes in the Santa Clara River systems. Symbols: +, species collected; S, species seen, but not collected.

10

Contributions in Science

No. 295

H

<1 on o — 1 (N m ^ in no c—

E— I ’ — M (N (N N (N (N N M

00

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CN CN co co co ro co

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• • + + CO • •

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snuoSnuaiop I

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kj

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— oj oi oj

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<N rj

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cn co co co co co

&

1978

Fishes of Santa Clara River

11

oo ON O I
m m ^

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"i- ^

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12

Contributions in Science

No. 295

(trout) and in favorable riffle and pool habitats of Sespe Creek. It was observed but
not collected above Lake Piru (stat. E). Lampetra tridentata is restricted to Sespe
Creek. One dead adult specimen was collected at station 31, one badly decomposed
adult was observed at station 32 and D. L. Soltz (personal communication) collected
several adults from near station 29 in June 1975. I also obtained two ammocetes
(LACM 35648) at station 30. Notemigonus crysoleucas, Dorosoma petenense, Cottus
asper and Micropterus salmoides were found mainly in the vicinity of Lake Piru.
Micropterus also was collected below Castaic Lake and in the Santa Clara River, 5
km downstream of the juncture with Castaic Creek (stat. 14). It also was observed
in private ponds along the upper reaches of the Santa Clara River (near stat. 19) but
not in the river. Cottus asper was found at stations 42, 44 and below Castaic Lake
at station 45 and was observed at station E, near Lake Piru. Dorosoma petenense and
Notemigonus crysoleucas were collected only in Lake Piru.

Eucyclogobius newberryi and Leptocottus armatus are species that frequently
occupy the mouths of rivers but fail to penetrate far inland. They were taken only at
station 1 . Other marine fishes that enter stream mouths might have been collected if
it had been possible to collect closer to the mouth of the river.

CONCLUSIONS AND DISCUSSION
Limitations of Data

To collect fishes by seining is at best an incomplete sampling method, although
it is the most practical one considering the large number of samples and the limited
resources available to make collections. The distributions here presented are based on
a series of collections made mostly during a short time span, obtained from a varying
number of seine hauls, generally from slow moving to still water because sampling
of sticklebacks was the primary objective. Seining is not ideal to obtain a complete
representation of the distribution of diverse fishes. Some species may have been
overlooked because special collecting techniques are required for capture. For in-
stance, Lampetra tridentata was found in Sespe Creek only by making a special trip
to look for adults during the spawning season. Rarer species, such as Lepomis cy-
anellus, might have been collected at more stations if collecting had been more in-
tense. Abundance or distribution of some species varies seasonally and these species
would have occurred at more stations had they been sampled year round. Thus, the
results of these collections are minimal measures of species diversity and distribution
in the Santa Clara River.

Native Fishes

Of the 15 fishes collected, Gasterosteus aculeatus, Eucyclogobius newberryi,
Leptocottus armatus, Salmo gairdneri and Lampetra tridentata are native. Gasterosteus
acculeatus was found in the headwaters of the Santa Clara River near Acton during
surveys for a Pacific railroad route in the middle 1800’s (Miller 1960). This form was
described as Gasterosteus williamsoni by Girard (1854) from “Williamson’s Pass,”
now known as Soledad Canyon (Miller 1960). Sticklebacks were present in the area

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Fishes of Santa Clara River

13

now occupied by the Santa Clara River system as early as the Pliocene (David 1945;
Bell 1973). Miller (1968) stated that Gasterosteus aculeatus was the only fish orig-
inally present in the Santa Clara River system. It appears to be native to most of the
system and in 1947 was collected in San Francisquito Canyon (CAS 20284) where
it may or may not be native. Two subspecies of G. aculeatus have been recognized
from this drainage (Miller and Hubbs 1969): G. A. williamsoni, mostly upstream of
Saugus, and G. a. microcephalus from the headwaters of Sespe Creek. Intergrades
between these two subspecies occupy much of the remainder of the drainage (Bell
1976).

Hubbs (1946) reported large and consistent runs of Salmo gairdneri into the Santa
Clara River. Sespe Creek between stations 31 and 35 is high gradient and consists of
riffles and pools. Although water temperauture is near the upper limits for Salmo
during the summer, they still are abundant. Thus, Salmo almost certainly is native to
Sespe Creek. Piru Creek and the Santa Clara River are much less suitable for S.
gairdneri. Rainbow trout are restricted to a few deep spots in the generally shallow
Piru Creek. California Department of Fish and Game records indicate that S. gairdneri
has been planted in Piru Creek since at least 1931, and these may be the original
source of trout observed at station E. Those observed at station 11 in the Santa Clara
River certainly had escaped from the Fillmore Hatchery just upstream of the station.
Although they were very abundant in the artificially cooled water flowing out of the
hatchery, they were absent 3 km downstream at station 10.

Eucyclogohius newberryi and Leptocottus armatus are euryhaline species that
have entered the Santa Clara River from the sea. There is no reason to believe that
they are introduced.

A fifth native species is Lampetra tridentata (placed in this genus by Hubbs
1971), the Pacific lamprey. Hubbs (1967) reported on three specimens from Sespe
Creek about 3 miles north of Fillmore (near stats. 30 and 31).

The native fishes of the Santa Clara River system have one thing in common,
they all are either anadromous or peripheral freshwater fishes. Peripheral freshwater
fishes are those restricted to freshwater immediately adjacent to the sea and the dis-
tributions of which are the result of dispersal through the sea (Darlington 1957),
whereas anadromous fishes spend a portion of their life cycle in marine water but
enter fresh water to spawn. Thus, all native fishes could have entered the Santa Clara
River system from the sea and there is no evidence that there has been any exchange
between the ichthyofauna of the Santa Clara River system and those of adjacent river
systems.

California Fishes Introduced
to the Santa Clara River System

Miller (1968) listed three fish species from California which have been introduced
to the Santa Clara River system. Rhinichthys osculus (Girard), the speckled dace, is
native to Santa Ana River system (Culver and Hubbs 1917), San Luis Obispo Creek
(based on Jordan 1894) and other coastal streams north of the latter. Miller (1968)
commented on the disjunct distribution of this species but concluded that it was in-
troduced to the Santa Clara River system. I did not collect this species there. Another

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native to the Santa Ana River system, Catostomus ( Pantosteus ) santaanae (Snyder
1908), apparently is introduced to the Santa Clara River system. Miller (1968:175)
concluded that the Santa Ana sucker is introduced based on the testimony of “An old
resident of that drainage. . . ” that a fish (i.e., Gasterosteus aculeatus) . . about
2 or 3 inches long, that swam in a jerky fashion and curled its tail when at rest.” was
the only species originally present. Catostomus santaanae had been introduced to the
Santa Clara River by 1934, when it also was found in Piru Creek (R. R. Miller per-
sonal communication). It was abundant in the Santa Clara River between Piru and
Fillmore, and in the lower reaches of Sespe Creek by 1940 (Hubbs et al. 1943). Hubbs
et al. (1943) reported another unidentified Catostomus of the subgenus Catostomus
and numerous hybrids between it and Catostomus santaanae collected between 1939
and 1942. Smith (1966) stated that C. santaanae from the Santa Clara River drainage
has features indicating introgression by a sucker of the subgenus Catostomus (i.e.,
papillae on the anterior face of the upper lip). The source of the second species of
Catostomus was unknown to Hubbs el al. (1943), but it is referred to the Owens
sucker, Catostomus fumeiventris Miller (1973) in his synonomy of that species. It
apparently was introduced by a release of Owens River water from the Los Angeles
Aqueduct. Owens suckers are endemic to the Owens River basin but have been in-
troduced to June Lake in Mono basin and the Santa Clara River drainage (Miller,
1973). I detected neither C . fumeiventris nor its hybrids and backcrosses with Catos-
tomus santaanae in my samples, but other recent surveys have reported them to be
present (A. W. Wells and J. S. Diana personal communication).

Gila orcutti is the third California species that has been introduced to the Santa
Clara River system. Miller (1968) also concluded that this species was introduced
because of the report that Gasterosteus aculeatus was the only native fish. Gila orcutti
is native to many streams from San Luis Rey River (Riverside County) north to Malibu
Creek (Los Angeles County) (Miller 1968). This species was first collected in the
Santa Clara River in 1934 (Miller 1968) when it also was found in Piru Creek (R. R.
Miller personal communication). Arroyo chubs were collected in San Francisquito
Canyon in 1947 (CAS 20253), and they are now the most widespread fish in the
system.

Cottus asper ranges south to Ventura County (Eddy 1957), and it has been col-
lected in the Ventura River (R. R. Miller personal communication) which enters the
Pacific Ocean just north of the Santa Clara River. Prickly sculpins were found only
in the vicinity of Lake Piru (stats. 42, 44, E) and Castaic Lake (stat. 45) but nowhere
else. R. R. Miller sampled Piru Creek several times since 1934 without collecting C.
asper (R. R. Miller personal communication). Both Lake Piru and Castaic Lake re-
ceive water from Pyramid Reservoir which receives its water from the Sacramento
and San Joaquin River drainages. C. asper occurs in the Sacramento River (Evermann
and Clark 1931) and this is probably the original source of all Cottus in the Santa
Clara River system.

Other Fishes Introduced
to the Santa Clara River System

Other fishes found in the Santa Clara River system are not native to California.
Pimephales promelas first was collected in California in 1950 and since has been

1978

Fishes of Santa Clara River

15

introduced to many waters (Shapovalov, Dill and Cordone 1959). Gambusia ajfnis
was introduced to California in 1922 and has become widespread for mosquito control
(Miller 1961). Lepomis cyanellus probably was introduced with bluegill (Lepomis
macrochirus Rafinesque) which it resembles when it is small (Evermann and Clark
1931). Ictalurus punctatus was first introduced to California in 1895 (Evermann and
Clark 1931) and has been introduced widely. Ictalurus me las (Rafinesque), the black
bullhead, was reported from near station 17 by J. N. Baskin (personal communica-
tion), but I did not collect it anywhere in the system.

Two fishes associated with Lake Piru probably were introduced there as forage
species for Micropterus salmoides, which was introduced for sport fishing. One, Do-
rosoma petenense, was introduced to California in 1953 because it is suited to the
warm fluctuating waters of reservoirs (Kimsey 1954). The other, Notemigonus cry-
soleucas, was reported in small creeks near San Diego by Evermann and Clark (1931)
and since has been introduced to many reservoirs.

Distribution Patterns

Threespine sticklebacks and Arroyo chubs are the most widespread fishes in the
Santa Clara River system. Native Gasterosteus aculeatus are found throughout the
drainage wherever there is slowly moving water, except in the smallest headwaters.
However, introduced Gila orcutti is more widespread, occurring farther upstream than
the native stickleback. Gila orcutti has dispersed well probably because it is native
to small coastal streams similar to the Santa Clara River system and can withstand
a stronger current than can Gasterosteus (J. N. Baskin personal communication). But
another coastal stream fish, Catostomus ( Pantosteus ) santaanae, apparently is not
sustained by the smaller flows (i.e., Arrastre Canyon, San Francisquito Canyon and
Todd Barranca) and has not penetrated above the low falls in Sespe Creek (stat. 32).

Three other introduced fishes, though widespread, have not penetrated higher
gradient tributaries. Gambusia ajfinis is present in the stream margins and pools of
low gradient areas but apparently has not become established in some of the steeper
tributaries. Mosquitofish have entered the upper Santa Clara River but have not spread
to its higher reaches. Pimephales promelas is restricted to low gradient portions of
the system except Piru Creek and below Castaic Lake. In Piru Creek, it was found
in Lake Piru and upstream (stat. E) and downstream (stats. 38, 39) of the lake. Fathead
minnows also occur at most stations in the Santa Clara River downstream of station
14 and only have penetrated a short distance up Sespe Creek (stat. 31). The distri-
bution of P. promelas indicates that it may have been introduced in imported water,
but unlike Cottus asper, successfully has dispersed. R. R. Miller (personal commu-
nication) over several years of collecting never obtained fathead minnows from Piru
Creek, supporting the view that it was introduced recently.

Native species such as Salmo gairdneri, Lampetra tridentata, Eucyclogobius
newberryi and Leptocottus armatus only are found in geographically restricted habi-
tats. Salmo gairdneri was found in cooler (stat. 11) and higher gradient (stats. 32 to
37) water (Table 1). Lampetra tridentata is restricted to the unique riffle and pool
habitat of Sespe Creek. Eucyclogobius newberryi and Leptocottus armatus were found
only at station 1 , near the sea.

Notemigonus crysoleucas, Dorosoma petenense and Micropterus salmoides ap-

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parently were introduced to Lake Piru and with the exception of the latter, are found
only there. Dorosoma petenense is a fragile fish (Kimsey 1954), which probably could
not survive in the small streams of the Santa Clara River system. Notemigonus cry-
soleucas also frequents quiet waters (Hubbs and Lagler 1952). Micropterus salmoides
favors sluggish waters also (Hubbs and Lagler 1952). Specimens at station 14 in the
Santa Clara River, which is typical stream habitat, may have been washed down from
Castaic Lake.

The distribution of Qottus asper can be easily explained. It was found just above
and below Lake Piru and below Castaic Lake and apparently was introduced in im-
ported water (see above). It may be such a recent arrival in the drainage that it has
not dispersed. High water temperature and other adverse habitat characteristics may
retard or limit its ultimate distribution.

Vulnerability of the Ichthyofauna

The Santa Clara River system includes diverse aquatic habitats ranging from high
gradient streams to sluggish, meandering streams with ponds and swampy areas.
These diverse habitats support at least 15 species of fishes which are threatened by
human activities. One problem is the increase of the human population centering
around Saugus. Human population growth probably will increase habitat destruction,
stream pollution, introduction of aquatic species, ground water exploitation and public
pressure to channelize streams in the drainage. Another problem is the storage of im-
ported water within the drainage.

Salmo gaireneri is probably immune to human disturbance because it is practi-
cally restricted to Los Padres National Forest. However, other fishes are more vul-
nerable because they are either found primarily in the Santa Clara River and lower
portions of tributaries that are, for the most part, outside of the national forest or pass
through this part of the drainage during spawning runs (i.e., Lampetra tridentata).
This part of the drainage, from about the town of Piru to the sea, is occupied by citrus
groves that seem to be compatible with the survival of fish populations. However, in
some places, water draining from the citrus orchards has left a residue of silt. It is
possible that fertilizers and pesticides also are being washed out of the orchards,
though there is no evidence.

While the citrus orchards along much of the Santa Clara River afford some pro-
tection for the fish fauna, urbanization rapidly is spreading near Saugus (between stats.
16 and 17) and in the Oxnard-Satacoy area (near stats. 1 and 2). Urban growth in. the
vicinity of Saugus already has produced some pressure to channelize parts of the Santa
Clara River drainage. On December 11, 1972, a hearing was held in Newhall, Cal-
ifornia where the U.S. Army Corps of Engineers proposed channelization of about
46 km (26.5 miles) of the Santa Clara River system. This proposal included chan-
nelization of about 18 km of river between Saugus and Lang, a generally dry stretch
of river. This proposed project would not result in the direct destruction of fish hab-
itats, but resultant increased runoff would have unpredictable consequences for hab-
itats downstream. If the proposed concrete-lined channel were constructed, continuous
water flow between the upper and lower portions of the Santa Clara River might result.
Continuous flow might allow movement of fishes across this area, allowing increased

1978

Fishes of Santa Clara River

17

introgression of the endangered subspecies, Gasterosteus aculeatus williamsoni (Miller
1972; California Fish and Game Code Sec. 2050 to 2055), by G. a. microcephalus
(unpublished data). G. a. williamsoni is a distinct subspecies divergent from other
Gasterosteus (Ross 1973; Bell 1976) and its introgression would constitute destruction
of an unusual subspecies that is protected by federal and state laws. Channelization
also might allow other fishes to invade the upper Santa Clara River.

Extension of channelization into the range of Gasterosteus aculeatus williamsoni
probably would result in its extinction in the Santa Clara River (J. N. Baskin personal
communication). An example of the effect of channelization can be seen in the lower
part of Santa Paula Creek. Santa Paula Creek at station C is comparable in gradient
(1.54%) to Todd Barranca at stations 26 and 27. Yet three or four species of fishes
(including Gasterosteus aculeatus ) live in the habitat of the latter two stations whereas
in the channelized portion of Santa Paula Creek there are no fishes.

Another threat to the fishes of the Santa Clara River is the introduction of or-
ganisms that adversely affect fishes. The probability of such introductions is increased
by greater human population density through release of bait and aquarium pets. Xen-
opus laevis, the African clawed frog, discovered by J. N. Baskin (personal commu-
nication) within the Santa Clara River drainage, probably was introduced by the re-
lease of pets and is considered to be a threat to fishes (St. Amant, Hoover and Stewart
1973). Aquatic organisms also may be introduced to the drainage with imported water.
Introgression of Catostomus santaanae by Catostomus fumeiventris apparently re-
sulted from the release of imported Owens River water. The storage of imported water
recently has increased within the Santa Clara River basin. Pyramid Reservoir began
to fill on January 6, 1972 and water immediately was released from it into Piru Creek.
Any fishes that have survived passage through pumping and power plants en route
to Pyramid Reservoir may colonize the Santa Clara River. I suggested above that
Cottus asper and Pimephales promelas may have been introduced by this means.
Colonization by other species may result in the introgression of Gasterosteus aculea-
tus, Gila orcutti and Catostomus santaanae, which are known to hybridize with con-
familial species in nature (Hagen 1967; Greenfield and Greenfield 1972; Hubbs et al.
1943). New introductions also may compete with fishes already present in the
drainage.

An estimate of the impact of urban growth and channelization may be obtained
by examining the ichthyofauna of the Los Angeles Basin. Culver and Hubbs (1917)
reported the presence of Lampetra tridentata, Gasterosteus aculeatus williamsoni,
Salmo gairdneri, Gila orcutti, Rhinichthys osculus and Catostomus santaanae in the
basin. Since 1917, most low gradient portions of the drainage have been urbanized,
most of the streams have been channelized and several fishes have been introduced.
Between 1929 and 1942 Gasterosteus aculeatus williamsoni, which previously was
abundant throughout the system (Culver and Hubbs 1971), became extinct in the basin
(Miller 1961). Miller (1961) attributed this extinction to the introduction of Gambusia
affinis. However, mosquitofish coexist with G. a. williamsoni in the Santa Clara
River, so its disappearance from the Los Angeles Basin probably was not caused
solely by the introduction of Gambusia. Miller (1961) also cited the deterioration of
surface flow as a contributory factor in the disappearance of G. a. williamsoni, and

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Contributions in Science

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this may have been critical. Recent efforts to collect Lampetra tridentata in the Los
Angeles Basin also have failed, and this species probably is extinct there (C. C. Swift
personal communication). The other four species listed by Culver and Hubbs (1917)
have persisted with much reduced ranges, although the presence of Catostomus san-
taanae is questionable. Thus, the condition of the fish fauna of the Los Angeles Basin
does not inspire optimism for the future of the fishes of the Santa Clara River system.

ACKNOWLEDGMENTS

C. Mahler, R. Putnam, D. L. Soltz and T. Webber assisted in field work. C.
C. Swift verified some identifications (for which I bear full responsibility) and pro-
vided stimulation and assistance. A. J. Cordone kindly allowed me to examine Cal-
ifornia Department of Fish and Game records. W. N. Eschmeyer allowed me to ex-
amine specimens under his care in the California Academy of Sciences. J. N. Baskin,
E. C. Olson and C. C. Swift offered constructive comments on the manuscript. I
thank B. M. Bell, my wife, who participated in most of the collecting and without
whom this research would have been impossible.

Personal communications from J. N. Baskin are based on his 1974 report, “Sur-
vey of the Unarmored Threespine Stickleback (i Gasterosteus aculeatus williamsoni )
in the Upper Santa Clara River Drainage.” (Unpublished final report for Bureau of
Sport Fisheries and Wildlife Contract No. 14- 16-001 -5387SE. iv + 67 p.) The per-
sonal communication from A. W. Wells and J. S. Diana is based on their 1975 report,
“Survey of the Freshwater Fishes and Their Habitats in the Coastal Drainages of
Southern California.” (Unpublished final report for California Department of Fish and
Game Contract No. AB-26. vi + 364 p.) I thank them for permitting me to cite their
unpublished data.

C. L. Hubbs and R. R. Miller were invaluable sources of unpublished infor-
mation, and I have depended extensively on their published and unpublished records.

This research was supported in part by the Theodore Roosevelt Memorial Fund;
University of California, Los Angeles Graduate Student Patent Fund No. 4-403828-
08613-3; Bureau of Sport Fisheries and Wildlife Contract No. 14-16-001-5387SE to
J. N. Baskin, Biological Sciences Department, California State Polytechnic Univer-
sity; the Section of Ichthyology, Natural History Museum of Los Angeles County,
and R. Bell, my mother.

LITERATURE CITED

Bailey, R. M., Chairman. 1970. A list of common and scientific names of fishes from the
United States and Canada, third edition. Amer. Fish. Soc. Spec. Publ. 6:1-150.

Bell, M. A. 1973. The Pliocene stickleback, Pungitius haynesi, a junior synonym of Gaster-
osteus aculeatus. Copeia 1973: 588-590.

— . 1976. The evolution of phenotypic diversity in threespine sticklebacks ( Gasterosteus

aculeatus). Ph. D. Dissertation, University of California, Los Angeles, xviii -I- 192 p.
Culver, G. B. and C. L. Hubbs. 1917. The fishes of the Santa Ana system of streams in
southern California. Lorquinia 1:82-83.

1978

Fishes of Santa Clara River

19

Darlington, P. J., Jr. 1957. Zoogeography: the geographical distribution of animals. John
Wiley and Sons, New York, xi + 675 p.

David, L. R. 1945. A Neogene stickleback from the Ridge Formation of California. Jour. Pa-
leont. 19:315-318.

Eddy, S. 1957. How to know the freshwater fishes. Brown Co. Publ. , Debuque, Iowa. 253 P.

Evermann, B. W. and H. W. Clark. 1931. A distributional list of the species of freshwater
fishes known to occur in California. Div. Fish Game California, Fish Bull. 35:1-67.

Girard, C. 1854. Descriptions of new fishes, collected by Dr. A. L. Heermann, naturalist
attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U.S.A.
Proc. Acad. Natur. Sci. Philadelphia 7:129-142.

Greenfield, D. W. and T. Greenfield. 1972. Introgressive hybridization between Gila or-
cutti and Hesperoleucus symmetricus (Pisces: Cyprinidae) in the Cuyama River basin, Cal-
ifornia: I. Meristics, morphometries and breeding. Copeia 1972:849-859.

Hagen, D. W. 1967. Isolating mechanisms in threespine sticklebacks ( Gasterosteus ). Jour. Fish.
Res. Bd. Canada 24:1637-1692.

Hubbs, C. L. 1946. Wandering of pink salmon and other salmonid fishes in southern California.
California Fish Game 32:81-86.

. 1967. Occurrence of the Pacific lamprey, Entosphenus tridentatus, off Baja California

and in streams of southern California with remarks on its nomenclature. Trans. San Diego
Soc. Natur. Hist. 14:303-311.

. 1971. Lampetra {Entosphenus) lathophaga, new species, the nonparasitic derivitive of

the Pacific lamprey. Trans. San Diego Soc. Natur. Hist. 16:125-163.

Hubbs, C. L... L. C. Hubbs and R. E. Johnson. 1943. Hybridization in nature between species
of catostomid fishes. Contrib. Lab. Vert. Biol., Univ. Michigan 22:1-76.

Hubbs, C. L. and K. F. Lagler. 1952. Fishes of the Great Lakes region. Cranbrook Inst.
Sci. Bull. No. 26, xi + 186 p.

Jordan, D. S. 1894. Notes on the fresh-water fishes of San Luis Obispo County, California.
Bull. U.S. Fish Comm. 14:141-142.

Kimsey, J. B. 1954. The introduction of redeye black bass and the threadfin shad into California.
California Fish Game 40:203-204.

Kimsey, J. B. and L. O. Fisk. 1960. Keys to the freshwater and anadromous fishes of Cali-
fornia. California Fish Game 46:453-479.

Miller, R. R. 1960. The type locality of Gasterosteus aculeatus williamsoni and its significance
in the taxonomy of California sticklebacks. Copeia 1960:348-350.

. 1961. Man and the changing fish fauna of the American southwest. Pap. Michigan

Acad. Sci., Arts, Lett. 46:365-404.

. 1968. Records of some native freshwater fishes transplanted into various waters of

California, Baja California and Nevada. California Fish Game 54:170-179.

. 1972. Threatened freshwater fishes of the United States. Trans. Amer. Fish. Soc.

101:239-252.

. 1973. Two new fishes, Gila hicolor snyderi and Catostomus fumeiventris, from the

Owens River basin, California. Occ. Pap. Mus. Zool. Univ. Michigan 667:1-19.

Miller, R. R. and C. L. Hubbs. 1969. Systematics of Gasterosteus aculeatus, with particular
reference to intergradation and introgression along the Pacific Coast of North America: a
commentary on a recent contribution. Copeia 1969:52-69.

Robins, C. R. and R. R. Miller. 1957. Classification, variation, and distribution of the scul-
pins, genus Cottus, inhabiting Pacific slope waters in California and southern Oregon, with
a key to the species. California Fish Game 43:213-233.

Ross, S. T. 1973. The systematics of Gasterosteus aculeatus (Pisces: Gasterosteidae) in central
and southern California. Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. 243:1-20.

Shapovalov, L. , W. A. Dill and A. J. Cordone. 1959. A revised checklist of the freshwater
and anadromous fishes of California. California Fish Game 45:159-180.

Smith, G. R. 1966. Distribution and evolution of the North American catostomid fishes of the

20

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No. 295

subgenus Pantosteus, genus Catostomus. Misc. Publ. Mus. Zool. Univ. Michigan 129:1-
132.

Snyder, J. O. 1908. Description of Pantosteus santa-anae, a new species of fish from the Santa
Ana River, California. Proc. U.S. Nat. Mus. 34:33-34.

St. Amant, J. A., F. G. Hoover and G. R. Stewart. 1973. African clawed frog, Xenopus
laevis (Daudin), established in California. California Fish Game 59:151-153.

Accepted for publication November 5, 1976.

507- 73

NUMBER 296
MAY 15, 1978

A NEW DEEP-SEA FISH FROM THE EASTERN NORTH PACIFIC
PSYCHROLUTES PHRICTUS (PISCES: COTTIDAE [PSYCHROLUTINAE] )

By David L. Stein and Carl E. Bond

Published by the NATURAL HISTORY MUSEUM
OF LOS ANGELES COUNTY
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Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

A NEW DEEP-SEA FISH FROM THE EASTERN NORTH PACIFIC
PSYCHROLUTES PHRICTUS (PISCES: COTTIDAE [PSYCHROLUTINAE])1

By David L. Stein2 and Carl E. Bond3

Abstract: Psychrolutes phrictus, new species, is described and compared with
its only congener P. paradoxus, from which it differs primarily in greater head length
(45.3 - 60.6% SL), larger adult size (over 500 mm), color pattern, and greater number
of pectoral fin rays (22-26). The systematic status of the subfamily Psychrolutinae
is discussed.

Psychrolutes phrictus is a very large benthic cottid, known between Monterey,
California, and northern Oregon at depths between 1006 m and 2800 m. Individuals
between 34 and 558 mm SL have been collected, all in otter trawls, beam trawls, or
benthic fish traps. We examined 19 preserved and three skeletonized specimens.

We concluded from analyses of stomach contents that P. phrictus is probably
an opportunistic feeder. Stomach contents included 24 different items; the most com-
mon of these were sea pens (three species), snails (two species), and crabs ( Chion -
oecetes sp.). Among other items found were ophiuroids, fishes, hermit crabs, octopod
beaks, and rocks. One specimen (309 mm SL) captured at 1097 m contained otoliths
of a large number of pelagic fishes. Capture of individuals of pelagic species by P.
phrictus probably depends upon those individuals swimming near the bottom; P.
phrictus does not appear to be capable of pelagic predation.

Since 1960, specimens of an undescribed, very large Psychrolutes have been
collected off the Pacific coast between Monterey, California, and northern Oregon.
The School of Oceanography of Oregon State University has collected individuals
between 34 and 469 mm SL on the continental slope off Oregon, at depths between
1026 and 2800 m. Acquisition of small and intermediate- sized specimens has made
a complete description of the species possible.

The family Psychrolutidae originally was characterized by lack of a spinous dor-
sal fin, presence of a suborbital stay, pseudobranchiae, thoracic ventrals, three and
a half gill arches, and naked skin (Gunther 1861:516). Jordan and Gilbert (1882:686)
indicated that P. paradoxus, the type species by monotypy, has a spinous dorsal fin
although it is buried in loose skin and flesh. Those authors and Gill (1889) believed
that differences between Psychrolutidae and Cottidae did not warrant a separate fam-
ily, and, therefore, included Psychrolutes in the latter family. Since then, some au-
thors (Jordan and Starks 1896; McCulloch 1926; Taranets 1941) maintained a separate
family designation for Psychrolutes; and others (Jordan and Evermann 1896; Jordan

Review Committee for this Contribution
J. D. McPhail
Joseph S. Nelson
Camm C. Swift

2School of Oceanography, Oregon State University, Corvallis, Oregon 97331.
department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331.

1

2

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No. 296

and Starks 1904; Hart 1973) included the genus in the Cottidae. We follow Gill
(1889), and consider the Psychrolutinae as a subfamily of the Cottidae.

METHODS

Counts and measurements follow Hubbs and Lagler (1964) except for counts
requiring dissection (pectoral fin, pelvic fin, gill rakers). These were made on the
right sides of the specimens. Pectoral and pelvic fin lengths are total (with dissection)
to base. Vertebral numbers were obtained from radiographs. Counts as given are the
mode, and in parentheses, the range. Morphometric ratios are given as the means,
with the range in parentheses; counts and ratios of the holotype are in brackets. Not
all measurements were made on all specimens because some individuals were badly
deformed during or subsequent to capture. The number of individuals examined (n)
follows each range. One specimen (OSUO 2040) was cleared and stained for com-
parison with P. paradoxus specimens that were treated similarly (OS 5300, 5301).

Stomach contents were examined without removal of stomachs from the speci-
mens. Many specimens had otoliths removed before our examination; the otoliths are
in the care of John E. Fitch, California Department of Fish and Game. Fitch also
removed and examined stomach contents of 16 fish. OSUO specimens were collected
by 3 m beam trawl (BMT) and 12 m otter trawl (OTB). Methods of collection for
other specimens are written out in materials examined.

MATERIALS

Specimens examined are on deposit at the National Museum of Natural History
(USNM), Auke Bay Biological Laboratory (ABBL), California Academy of Sciences
(CAS), Natural History Museum of Los Angeles County (LACM), Department of
Fisheries and Wildlife, Oregon State University (OS), School of Oceanography, Or-
egon State University (OSUO), and University of British Columbia (BC).

ACKNOWLEDGMENTS

We thank N. J. Wilimovsky, (BC) for very kindly relinquishing his priority in
working on these fishes, assisting us in our description, and lending specimens ofP.
paradoxus.

For loan of specimens, we thank T. Iwamoto and W. Eschmeyer (CAS), E. Lu
Hall (ABBL), and R. Lavenberg (LACM) who also greatly helped to facilitate this
work. A. Welander, University of Washington, donated specimens of P. paradoxus
(OS 5300, 5301). J. S. Nelson, University of Alberta, freely gave very useful advice
and encouraged us.

Joanne Laroche (OSUO) cleared and stained specimens for us and supplied in-
formation on cottid development.

John Fitch, California Dept, of Fish and Game, supplied information on stomach

1978

New Deep-Sea Fish from Eastern North Pacific

3

contents. The manuscript was reviewed by W. G. Pearcy (OSUO), B. J. Verts (OS),
and J. S. Nelson. Bond was supported by Oregon Agricultural Experiment Station.

This work was supported by ERDA Contract E(45-l) - 2227 Task Agreement
12 (RLO-2227-T 12-68).

Psychrolutes phrictus new species
Figures 1 and 2.

Holotype: USNM 216253 (ex OSUO 1839), 1 (383 mm SL, 9), OTB 92, 44°44.3'N, 125°41.3'W,
2800 m, 24 October 1965.

Paratypes: CAS 32580, 1 (478 mm SL, 9), sablefish trap, 36°44.5'N, 122°04.5'W, 600 fm
(1097 m), 22 April 1975; LACM 35230-1, 3 (495, 522, 558 mm SL, 9 9 9), otter trawl,
40°37.3'N, 124°43.0'W, 600 fm, (1097 m), 29 November 1974; LACM 35232-1, 1 (519
mm SL, 9), otter trawl, 40°37.3'N, 124°43.0'W, 600 fm (1097 m), 14 November 1974;
LACM 34338-1, 1 (380 mm SL, 9), otter trawl?, 40°45.3'N, 124°47.4'W, 700 fm (1280
m), 13 February 1974; LACM 35234-1, 1 (501 mm SL, 9), otter trawl, 40°45.7'N,
124°44.0'W, 600 fm (1097 m), 3 April 1975; LACM 34185-1, 2 (475, 522 mm SL, ? 9),
otter trawl, 40°45.8'N, 124°43.3'W, 550 fm (1006 m), 26 August 1974; LACM 35561-1,

1 (298 mm SL, 6), otter trawl? approx. 40°46.0'N, 124°44.0'W, 600 fm (1097 m), 8
August 1972; LACM 30807-4, 1 (508 mm SL, 9), otter trawl, 41°21.6'N, 124°56.9'W,
725 fm (1326 m), 25 September 1969; OSUO 2057, 1 (440 mm SL, 9), BMT 419,
42°51.9'N, 124°59.5'W, 1026 m, 17 September 1974; OSUO 2221, 1 (148 mm SL, 9),
OTB 500, 43°22.0'N, 125°09.9'W, 1600 m, 4 April 1973; OSUO 2029, 1 (469 mm SL,
9), OTB 634, 43°32.0'N, 125°13.0'W, 1580 m, 7 July 1974; OSUO 1524, 1 (399 mm
SL, 9), OTB 64, 44°32.5'N, 125°24.0'W, 2800 m, 9 April 1965; OSUO 2020-2021, 2
(34, 49 mm SL, immatures), OTB 208, 44°36.0'N, 125°10.8'W, 1390 m, 30 October 1967;
OSUO 2040, 1 (37 mm SL, immature), OTB 205, 45°51.4'N, 125°15.2'W, 1600 m, 25
October 1967 (cleared and stained).

Additional non-type material: LACM 35771-1, 1 (555 mm SL, 9), otter trawl, 40°44.3'N,
124°42.3'W, 510-565 fm (933-1051 m), 30 November 1972 (skeleton); LACM 35770-1,

2 (538, 468 mm SL, 9 9), otter trawl, 40°40.5'N, 124°44.8'W, 580-600 fm (1061-1097
m), 27 February 1973 (skeletons).

Other Collections. — Four specimens held at the University of British Columbia
were not examined by us. These were collected off Oregon in 1963.

Comparative Material. — Psychrolutes paradoxus Gunther 1861: OS 5301, 1 (35
mm SL, ?) Lopez Is., San Juan Islands, Washington, 10 July 1963 (cleared and
stained); BC 53-260, 6 (27-29 mm SL, ?), Friday Harbor, San Juan Islands, Wash-
ington, July 1950; ABBL 72-74, 1 (42 mm SL, 9), 5 (about 25 mm SL, ?), southeast
shore of Favorite Channel, vicinity of beach between Pt. Louisa and Pt. Lena, near
Juneau, Alaska, 10 November 1972; ABBL 64-755, 7 (45-48 mm SL, ?), Sta. 755,
northeast of Afognak Is., Gulf of Alaska, 58°26'N, 151°5UW, 13 August 1963; OS
5300, 1 (37 mm SL, ?), Kodiak Is., Gulf of Alaska, 15 June 1970 (cleared and
stained).

Psychrolutes zebra Bean 1891: Holotype, USNM 45364, sex undetermined,
“Albatross” sta. 2848, 55°10'N, 160°18'W, 110 fm (201 m), 31 July 1888.

Diagnosis. — A Psychrolutes differing from its only congener in the following:
Head large, its length 45.3-60.6% SL; gill rakers on first arch 9-13; pectoral fin rays
22-26. Head with small cirri, especially above eyes and on snout and lower jaw;

4

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No. 296

'i

Figure 1. Psychrolutes phrictus. Holotype; 383 mm SL; USNM 216253.

1978

New Deep-Sea Fish from Eastern North Pacific

5

Figure 2. Photograph of Psychrolutes phrictus. Holotype; 383 mm SL; USNM 216253.

6

Contributions in Science

No. 296

scattered cirri on body. Prickles as well as cirri present on heads and bodies of in-
dividuals less than 50 mm SL. Color grayish or blackish, sometimes mottled with
white, ventral surface often white, but no distinct banding or sharply defined pattern.

Description. — Dorsal fin rays VIII, 20 [VIII, 20] (VII-IX, 19-20) (n=19), anal
fin rays 13 [13] (12-14) (n=19), pectoral fin rays 24 [23] (22-26) (n=19), pelvic fin
rays I, 3 (n=19), principal caudal rays about 13 [type not dissected]. Gill rakers on
first arch 11 [12] (9-13) (n=19). Lateral line pores 12 [12] (12-14) (n=16). Vertebrae
[33] (33-35) (n=4).

Head length 48.9 [50.7] (43.5-60.6% SL) (n=14);eye 1. 12.8 [9.1] (8.6-24.3%
head 1.) (n=14), fleshy interorbital width 35.6 [33.2] (24.4-46.4% head 1.) (n=9),
pectoral 1. 55.3 [55.4] (44.9-62.3% head 1.) (n=14), pelvic 1. 26.8 [24.1] (23.3-
34.7% head 1.) (n=14), snout-anus 109.8 [105.7] (93.8-132.2% head 1.) (n=15).

Allometry is evident over the size range of individuals examined. Small speci-
mens have longer heads, larger exposed portions of the eye, and an apparently nar-
rower interorbital space than large specimens.

Head unusually large, broad, and flattened. In smaller individuals (less than 50
mm SL), depth at occiput about equal to head width; in larger specimens, depth at
occiput about 66% of head width. Nostrils two, the anterior with a very distinct tube,
about on a horizontal plane with suborbital stay; the posterior on a horizontal line
through pupil of eye, distance anterior to eye about equal to eye diameter. Interorbital
space broad, slightly convex. Orbits large, orbital rims very poorly ossified, especially
in large individuals, dissection required for their accurate measurement. Eyes rela-
tively large, but exposed portion reduced in specimens greater than 150 mm SL.
Mouth broadly curved, slightly oblique, lips fleshy; in individuals greater than 150
mm SL, lower jaw distinctly included. Premaxillaries not reaching fleshy rictus; a
wide, distinct gap at medial juncture, a narrower gap present at mandibular symphysis.
Premaxillary teeth sharp, stout, small and recurved, posteriorly arranged in irregular
oblique rows, especially in small specimens, forming band about four teeth wide in
individuals less than 50 mm SL, up to eight teeth wide in large (400 mm SL) spec-
imens. Tooth band becoming uniserial posteriorly in small individuals, in larger ones
not narrowing much, not uniserial posteriorly.

Cephalic sensory canals well developed, anterior sensory pores large, distinct,
posteriormost pores smaller, difficult to distinguish. Five pores in a series above max-
illary, eight in the preoperculomandibular series, one nasal pore close and lateral to
anterior nostril. Anterior mandibular pores well separated.

Small cirri present on head as slender, short, sharply pointed filaments, usually
single, although some on mandible are paired. Mandibular cirri form a line behind
lips although no distinct pattern present; other cirri scattered over ventral surface of
mandible. Cirri on upper part of head located between maxillary pores, on snout,
interorbital region, and posterior to upper orbits. A line of widely separated cirri pres-
ent along upper orbital margin. Other cirri scattered widely over entire head and body.

Opercular ossification poor, musculature weak, covered with gelatinous layer.
Opercular opening long, extending from far above pectoral fin base to just below it.

Body behind opercular flaps tapering rapidly to caudal fin. Pectoral fins broad,
well developed, their bases oblique, rays (except uppermost) evenly graduated in

1978

New Deep-Sea Fish from Eastern North Pacific

7

length to shortest ray at anterior (ventral) point of pectoral base. In specimens above
150 mm SL, approximately the 10 lowest rays with distinct fleshy pads on outer tips;
in largest specimens, pads very thick, tough, pale colored.

Pelvic fins with one spine and three soft rays. Length of pelvic fin spine about
50% of total fin length, the first (outer) ray almost equal to inner two rays which are
about equal in length. Basal half of fin hidden beneath body skin, apparently not very
erectile. Distal half of fin free, covered with thick skin, only tips of rays distinguish-
able without dissection. Pelvic bases narrowly separated. Pelvic fins relatively longer
in small specimens.

Dorsal fin with spinous and soft-rayed portions. Anterior spinous portion deeply
buried in loose skin and gelatinous tissue, its origin anterior to upper end of opercular
opening, spines well developed, stout, although flexible. Spinous portion externally
marked by short free filaments above most spines. A shallow notch present between
spinous and soft-rayed fin sections. Soft dorsal fin rays becoming gradually longer
posteriorly, fin becoming high and distinctly lobate at its end. Anal fin well developed,
of soft rays only, its origin below anterior dorsal fin rays.

Anus distant from origin of anal fin, anterior to a vertical through pectoral tips.
Body covered with thick, tough skin, naked, except in specimens less than 50 mm
SL, which have simple, sharp, slender prickles distributed over body except head and
parts of fins, especially dense around anus. Larger individuals with a thick gelatinous
layer between skin and musculature, often with distinct, yellowish fat deposits in
concavities of muscles and bones. In specimens greater than 300 mm SL, skin ex-
tensively marked with pale lines, circles, and other evidences of injury. Largest spec-
imens (above 400 mm SL) with very distinct areas irregularly covered with pale rings,
often overlapping in great numbers. We believe the rings are sucker marks of cephalo-
pods, a known food item. Patterns of rings fit the disposition of suckers on octopod
arms.

Lateral line consisting of 12-14 open pores, usually well spaced, but occasionally
with two pores close together. Posteriormost two pores located on caudal base or on
caudal fin itself. Anteriormost lateral line pore reduced, above gill opening.

Color of skin grayish or blackish dorsally, often pale ventrally, with indistinct
mottling especially on head. Mouth pale, opercular cavity pale, peritoneum pale,
stomach pale.

Food Habits. — Although the data obtained through examination of stomach con-
tents do not warrant statistical analysis, some speculations are possible about the feed-
ing habits of P. phrictus. This species apparently has a varied diet; a total of 24
different food items was found in the 25 stomachs examined. The most frequent food
items were sea pens ( Stylatula sp., Balticina ? sp., Funiculina sp.) in 12 fish; crabs
( Chionoecetes sp.) in 15 fish; and snails ( Buccinium sp., unidentified sp.) in 13 spec-
imens. Other food items were Sebastolobus alascanus, Sebastes sp., octopod beaks
(in 4 individuals), ophiuroids, a plastic bag, rocks, hermit crabs {Pagurus sp., Par-
apagurus sp.), and possible anemone, sea cucumber, and crinoid remains. One fish,
LACM 35561, 309 mm SL, contained a large number of otoliths of Nansenia sp. and
Lestidium sp. in its stomach, plus otoliths of Sebastes sp. and T etragonurus sp. Except
for Sebastes, all are known to be pelagic. Although this individual was the smallest

Contributions in Science

No. 296

with stomach contents, we have no other reason to believe it was pelagic. Several of
the OSUO specimens were smaller, and were captured with bottom trawls. Because
the anatomy ofP. phrictus is typical of a demersal cottid, we believe that the pelagic
species ingested must have become available by swimming near the bottom. The pos-
sibility that individuals of P . phrictus as small as 300 mm are pelagic is unlikely
because all specimens, including those smaller than 300 mm, were taken by bottom
trawl.

The presence of octopod beaks in the stomach contents is clear evidence that P.
phrictus sometimes eats octopods. This may account for the scars on the skin of the
head in large specimens. Some of the beaks are large: specimen OSUO 2029, 469
mm SL, contained a beak from an octopod estimated (by the method of Clarke 1962)
to weigh about 1 kg.

Distribution. — Psychrolutes phrictus occurs along the middle and lower conti-
nental slope from Monterey (1097 m) to Eureka (1006-1326 m), and off Oregon
(1026-2800 m). Its meridional distribution may be wider, especially towards the north
where there seems to be no hydrographic or geologic features that might act as
barriers.

Males and females may be distributed differently. Of the 25 specimens of known
sex, two are males and only one of the 19 fish we examined was a male (LACM
35561-1, 298 mm SL). The other male specimen (Calif. Fish and Game, Eureka Lab.
No. 68, 475 mm SL) was not seen by us. Perhaps males are segregated in rocky areas
where trawling is difficult.

Etymology . — From typiKrocr (phriktos), Greek, “causing one to shudder,” from
the grotesque appearance of the species.

Relationships . — Psychrolutes phrictus is provisionally thought to be most closely
related to P. paradoxus Gunther 1861. It is distinctly different in the following re-
spects: the head is relatively longer, especially in small specimens; those of 37 to 58
mm have head lengths of 49.6 to 60.6% with the overall range 45.3-60.6% of SL vs.
P. paradoxus 40.2-44.4%; it has a larger pectoral fin of 25 (22-26) rays vs. P. par-
adoxus 21 (20-22) rays. It has cirri whereas P. paradoxus has large numbers of stout,
blunt, papillae over its entire body, including outer surfaces of the fins; small spec-
imens have many sharp exposed prickles whereas large P. paradoxus have a few in
two ventrolateral rows beneath the skin. Its two inner pelvic fin rays are about equal,
with the outer ray slightly shorter; P. paradoxus has pelvic fin rays evenly graduated
in length, the outer shortest. Body color is generally evenly grayish, blackish, or
mottled, whereas that of P. paradoxus is brown or black, in sharply defined bars or
blotches on a light background, in a consistent pattern. Maximum size is at least 558
mm, whereas that of P. paradoxus is 58 mm (Hart 1973:534). Depth distribution
differs: P. phrictus occurs between 1006-2800 m, but P. paradoxus is found between
55-220 m (Hart 1973:534).

Discussion. — This is the only other species beside P. paradoxus now referred
to Psychrolutes; two other species were described but later removed from the genus
or synonymized. Psychrolutes latus Hutton was redescribed by Gunther (1876) as the
type of a new genus, Neophrynichthys; Psychrolutes zebra Bean (1891) was synon-
ymized with P. paradoxus by Jordan and Evermann (1896:2027), with whom we
agree.

1978

New Deep-Sea Fish from Eastern North Pacific

9

The relationships of Psychrolutes and closely related genera (e.g. Neophrynich-
thys Gunther and Cottunculoides Barnard) are not well known; the characters sepa-
rating them are unclear. If future investigations result in merging of these genera,
Psychrolutes will be the senior synonym. We have placed the new species into Psy-
chrolutes because it agrees with the generic description of the following characters:
Lack of spines on the head, lack of opercular spines, pelvic fins I, 3, lack of scales
on the body, seven branchiostegal rays, vomerine and palatine teeth absent, and dorsal
fin spines completely buried in flesh or gelatinous tissue. Comparison ofP. paradoxus
and P. phrictus using the above characters leaves little doubt that they are closely
related.

LITERATURE CITED

Bean, T. H. 1891. Scientific results of explorations by the U.S. Fish Commission Steamer
“Albatross”. No. XL New fishes collected off the coast of Alaska and the adjacent region
southward. Proc. U.S. Natl. Mus. 13:37-45.

Clarke, M. R. 1962. The identification of cephalopod “beaks” and the relationship between
beak size and total body weight. Bull. British Mus. (Nat. Hist.) Zoology. 8(10):421— 485.
Gill, T. 1889. On the Psychrolutidae of Gunther. Proc. U.S. Natl. Mus. 11:321-327.
Gunther, A. 1861. Catalog of the Acanthopterygian Fishes in the Collection of the British
Museum. 3:1-586.

1876. Remarks on fishes, with descriptions of new species in the British Museum,

chiefly from southern seas. Ann. Nat. Hist. ser. 4. 17:389-402.

Hart, J. L. 1973. Pacific Fishes of Canada. Fish. Res. Bd. Canada Bull. 180:1-740.

Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great Lakes Region. Revised edition.

University of Michigan Press, Ann Arbor. 213 pp.

Jordan, D. S. and B. W. Evermann. 1896. The Fishes of North and Middle America. Bull.
U.S. Natl. Mus. 47:1-3313.

and C. H. Gilbert. 1882. Synopsis of the Fishes of North America. Smithsonian

Misc. Coll. 24:1-1074.

and E. C. Starks. 1896. The fishes of Puget Sound. Proc. Calif. Acad. Sci. ser.

2, 5:785-855.

1904. A review of the Cottidae or sculpins found in the waters of Japan. Proc. U.S.

Natl. Mus. 27( 1 358):23 1—335.

McCulloch, A. R. 1926. Report on some fishes obtained by the F.I.S. “Endeavour” on the
coasts of Queensland, New South Wales, Victoria, Tasmania, South and south-western
Australia. Biol. Res. F.I.S. “Endeavour”, 5(4): 157-216.

Tara nets, A. Y. 1941. K klassifikatsii i proiskhozhdenif bychkov semeystva Cottidae. Izvestiya
Akad. Nauk SSSR. Otd. Biol., 3:427-447. On the classification and origin of the family
Cottidae. Transl. N. J. Wilimovsky and E. Lanz. 1959. Museum Contr. 5, Inst. Fish.,
Univ. Brit. Columbia, Vancouver, Canada.

Accepted for publication January 9, 1977.

507. 73

NUMBER 297
MAY 22, 1978

A NEW GENUS AND SPECIES OF CERATIOID ANGLERFISH FROM THE
NORTH PACIFIC OCEAN WITH A REVIEW OF THE ALLIED GENERA
CTENOCHIRICHTHYS, CHIROPHRYNE AND LEPTACANTHICHTHYS

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

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A NEW GENUS AND SPECIES OF CERATIOID ANGLERFISH FROM THE
NORTH PACIFIC OCEAN WITH A REVIEW OF THE ALLIED GENERA
CT ENO CH1RICHT HYS , CH1ROPHRYNE AND LEPTACANTHICHTHYS 1

By Theodore W. Pietsch2

Abstract: A new genus and species of deep-sea anglerfish is described on the
basis of two specimens collected from the north Pacific Ocean. The new form is most
strikingly characterized by having an unusually elongate pectoral-fin lobe that bears
the fin rays along its dorsal margin, a unique and derived condition shared with three
other oneirodid genera .Ctenochirichthys, Chirophryne, and Leptacanthichthys . The
new form is described and compared osteologically with its nearest allies. It differs
in having an extremely short snout, strongly convex frontal bones, and an oblique
jaw suspension, appearing to be intermediate in many ways between more generalized
oneirodids and the other three “long-pectoraled” forms. It is probably most closely
related phylogenetically to Ctenochirichthys. Chirophryne is perhaps intermediate
between the new genus and Ctenochirichthys on one hand, and Leptacanthichthys on
the other. A key to the “long-pectoraled” genera is provided.

INTRODUCTION

Among the previously described genera of the deep-sea anglerfish family Onei-
rodidae are three forms unique among the Ceratioidei in having an unusually elongate
pectoral-fin lobe that bears the fin rays along its dorsal margin. These genera are
Leptacanthichthys, Chirophryne, and Ctenochirichthys. Leptacanthichthys was orig-
inally described by Regan and Trewavas (1932) as one of five subgenera of the genus
Dolopichthys Garman (1899) to include a single species, D. gracilispinis Regan
(1925), based on two metamorphosed female specimens. Since that time the name
Leptacanthichthys has been given generic status (Bertelsen 1951:94). Pietsch (1974)
described the osteology of the genus and discussed its phylogenetic relationships with
other oneirodid genera. More recently, Pietsch (1976) reported a case of sexual par-
asitism in Leptacanthichthys, the first known occurrence of this peculiar mode of
reproduction in the family Oneirodidae and the first record of a male for the genus.

Chirophryne and Ctenochirichthys were introduced by Regan and Trewavas
(1932) as monotypic genera: Chirophryne xenolophus, described from a single met-
amorphosed female, and Ctenochirichthys longimanus , based on two metamorphosed

Review Committee for this Contribution
E. Bertelsen
Robert J. Lavenberg
Richard H. Rosenblatt
Camm C. Swift

2Research Associate in Ichthyology, Natural History Museum, Los Angeles, California 90007,
and Department of Biology, California State University, Long Beach, California 90840

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No. 297

females. Since the original publication a second female ofC. xenolophus was reported
by Pietsch (1974:31, fig. 58). Two additional larval males and an adolescent male of
C. longimanus were reported by Bertelsen (1951:95) and Beebe and Crane (1947:166),
respectively.

Recently, a fourth “long-pectoraled” oneirodid, representing an undescribed
genus and species, was discovered among the ceratioids in the collections of the In-
stitute of Oceanography of the Academy of Sciences of the USSR, Moscow, and the
Scripps Institution of Oceanography in La Jolla. This new form is described and com-
pared with its nearest allies below.

METHODS AND MATERIALS

Standard lengths (SL) are used throughout. Measurements were taken on the left
side whenever possible and rounded to the nearest 0.5 mm. To insure accurate fin-
ray counts skin was removed from the pectoral fins and incisions were made in the
skin to reveal the rays of the dorsal and anal fins. Sockets, indicating missing teeth
in the jaws and on the vomer, were included in total tooth counts. Jaw tooth counts
are the sum of left and right sides. Head length is the distance from the tip of the
upper jaw to the posteriormost margin of the preoperculum; head depth is the distance
from the tip of the sphenotic spine to the base of the quadrate spine; lower jaw length
is the distance from the symphysial spine to the posteriormost margin of the articular;
illicium length is the distance from the articulation of the pterygiophore of the illicium
and the illicial bone to the dorsal surface of the escal bulb, excluding escal appen-
dages. Terminology used in describing the various parts of the angling apparatus fol-
lows Bradbury (1967). Definitions of terms used for the different stages of devel-
opment follow Bertelsen (1951:10-11). Study material is deposited in the following
institutions: BMNH — British Museum (Natural History), London; IOS — Institute of
Oceanographic Sciences, Surrey, England (formerly the National Institute of Ocean-
ography); LACM — Natural History Museum of Los Angeles County; MCZ — Mu-
seum of Comparative Zoology, Harvard University; ROM— Royal Ontario Museum,
Toronto; SIO — Scripps Institution of Oceanography, La Jolla; CAS-SU — Stanford
University (collections now housed at the California Academy of Sciences, San Fran-
cisco [CAS]); ZMUC — Zoological Museum, University of Copenhagen.

OSTEOLOGY

The osteology of Leptacanthichthys previously has been described in detail and
compared with that of other oneirodid genera (Pietsch 1974). Material sufficient for
a thorough osteological study of Ctenochirichthys and Chirophryne is unavailable; a
few comparative aspects however, are discussed based on superficial dissection. An
osteological preparation of one of the two known specimens of Puck (SIO H52-363),
made using the trypsin digestion technique of Taylor (1967), forms the basis for the
following description.

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N. Pacific Ocean Ceratioid Anglerfish

3

Cranium. — The T-shaped supraethmoid of Puck has a long ventromedial process
that forms, together with the lateral ethmoids and relatively thin, dorsally concave
ethmoid cartilage, large, roughly circular, nasal foramina (Fig. 1). This is similar to
the condition found in Oneirodes and Danaphryne (Pietsch 1974:16, fig. 22). Like
Danaphryne, but again in contrast to Leptacanthichthys, the illicial trough of Puck
is deep and narrow resulting from a lateral compression of the anterior portion of the
skull, rather than a medial, dorsoventral depression of the ethmoid region. The width
of the ethmoid cartilage and underlying vomer of Puck is greater than the distance
between the anterolateral tips of the lateral ethmoids (Fig. 1).

The frontal bones of Puck (Fig. 2) are similar to those of Oneirodes and Dan-
aphryne, having a strongly convex dorsal margin and occupying a relatively posterior
position on the skull. The frontals of Leptacanthichthys are considerably longer, have
a nearly linear dorsal margin, and occupy a more anterior position, overhanging and
extending past the anterior limits of the ethmoid cartilage and vomer (Pietsch 1974:18-
19, figs. 2, 28-31, 34). Ctenochirichthys and Chirophryne have moderately convex
frontals (Figs. 11, 14).

The sphenotics of Puck, Chirophryne, and Leptacanthichthys are large, forming

Figure 1. Anterior view of anterior half of cranium of Puck pinnata, SIO H52-363, 66.0 mm.

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Figure 2. Lateral view of cranium of Puck pinnata, SIO H52-363, 66.0 mm.

an elongate spine (Figs. 2, 9, 14, 16; Pietsch 1974:19, fig. 34). The sphenotic spines
of Ctenochirichthys are short, in some specimens not piercing the skin (Fig. 11).

Mandibular, palatine, and hyoid arches. - Puck differs from other “long-pector-
aled” oneirodids in the angle of the jaw suspension. The suspensorium of Puck is
strongly oblique in a posterior direction, resulting in a more posterior position for the
lower jaw. In contrast, the angle of the jaw suspension in Ctenochirichthys, Chiro-
phryne, and Leptacanthichthys is nearly vertical; the lower jaw extends forward
slightly beyond the premaxillaries.

Puck and Ctenochirichthys are similar in that the distal ends of the upper jaw
bones are held in a forward position on the dentary by a relatively short anterior
maxillomandibular ligament, resulting in a short gape that terminates anterior to the
eye (Fig. 3A, B, 9, 11). Although, the anterior maxillomandibular ligament of Cten-
ochirichthys is slightly longer than that of Puck, it originates much further forward
on the dentary. For this reason, the length of the gape of Ctenochirichthys is slightly
shorter than that of Puck. The anterior maxillomandibular ligament of Chirophryne
and Leptacanthichthys is considerably longer than that of Puck and Ctenochirichthys,
the upper jaw bones are more horizontal in position, and the gape of the mouth extends
past the level of the eye (Figs. 3C, D, 14, 16).

In apparent correlation with this difference in jaw mechanism, the suspensoria
(as well as the opercular apparatus and pectoral girdles) of Puck and Ctenochirichthys
flare out laterally so that viewed head-on, the crania of these genera are considerably
more narrow than the distance between the posterior ends of the lower jaw (Fig. 13 A,
B). Thus, the mouths of these two genera are short, yet wide, relative to other onei-

1978

N. Pacific Ocean Ceratioid Anglerfish

5

C D

Figure 3. Lateral views of jaws with anterior maxillomandibular ligaments indicated by stip-
pling: A. Puck pinnata, holotype, LACM 34276-1, 46.0 mm; B. Ctenochirichthys longimanus,
lectotype, ZMUC P9297, 12.5 mm; C. Chirophryne xenolophus, SIO 70-306, 22.0 mm; D.
Leptacanthichthys gracilispinis, LACM 33625-2, 56.0 mm.

rodids. In contrast, the crania of Chirophryne and Leptacanthichthys are equal in
width, or slightly wider than the distance between the posterior ends of the lower jaw
(Fig. 13C, D); their mouths are consequently longer, yet more narrow.

Among oneirodids, differences in the size and shape of the bones of the lower
jaw are correlated with differences in the length and depth of the cranium (Pietsch
1974:20). Puck has a relatively short, deep cranium, and consequently, like Oneirodes
and Danaphryne, has short, deep dentaries and articular bones (Fig. 4; Pietsch 1974:

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No. 297

Figure 4. Lateral view of left side of lower jaw, suspensorium and opercular apparatus of Puck
pinnata, SIO H52-363, 66.0 mm.

9, 20-21, figs. 8, 40). Leptacanthichthys, on the other hand, is a long-jawed genus
with a more elongate, narrow cranium (Pietsch 1974: 20-21, fig. 42).

Variation in the length and width of the palatine arch is also associated with the
length and depth of the cranium. Puck, Oneirodes, and Danaphryne have short, broad
mesopterygoid, ectopterygoid and palatine bones as compared to Leptacanthichthys
in which these bones are considerably more elongate and narrow (Fig. 4; Pietsch 1974:
9-10, 21, figs. 8, 40, 42).

At the articulation of the lower jaw and suspensorium, the articular and quadrate
bones form spines (the mandibular and quadrate spines, respectively), which vary in
relative length among oneirodids. The mandibular spine is considerably shorter than
the quadrate spine in Puck and Chirophryne (Figs. 5A, 14), but, in contrast to all
other oneirodids, the mandibular spine is considerably longer than the quadrate spine
in Leptacanthichthys (Fig. 5B). Both spines are rudimentary in Ctenochirichthys (Fig.
11, Table 2).

In all oneirodids the jaw teeth are slender, recurved and depressible, large and
small ones intermixed in both jaws. The considerably longer gape of Leptacanthich-
thys provides space for a greater number of upper and lower jaw teeth than are found
in Puck and Ctenochirichthys. Chirophryne, known from only two small, adolescent
females (11.0 and 22.0 mm), doubtless has a greater number of jaw teeth in larger
specimens (Table 2).

Opercular apparatus. — The shape of the subopercular bone varies considerably

1978

N. Pacific Ocean Ceratioid Anglerfish

7

Figure 5. Ventrolateral views of posterior part of lower jaw, left side, showing arrangement
of quadrate and mandibular spines: A. Puck pinnata, holotype, LACM 34276-1, 46.0 mm; B.
Leptacanthichthys gracilispinis, LACM 33625-2, 56.0 mm.

between genera and is a diagnostic feature of many oneirodids. The subopercula of
Puck and Ctenochirichthys are elongate and slender throughout their length, the upper
end tapering to a point (Fig. 6A-C; Bertelsen 1951:94, fig. 51). Those of Leptacan-
thichthys and the two known specimens of Chirophryne (11.0 and 22.0 mm) are short
and broad, the upper end rounded to bluntly pointed (Fig. 6D-G; Bertelsen 1951:94,
figs. 49, 50; subopercula of other oneirodids are figured by Pietsch 1974).

Branchial arches. — The branchial arches of Puck are similar to those of Dana-
phryne and Leptacanthichthys (Pietsch 1974:25). Pharyngobranchials II and III are
well developed and bear eight and ten long, slender teeth, respectively (Fig. 7; Pietsch
1974:25, fig. 51A, C).

Dorsal and anal fins. — Puck, Chirophryne and most specimens of Leptacanthich-
thys have five dorsal rays; Ctenochirichthys has six or seven. Puck, Chirophryne and
most individuals of Ctenochirichthys have four anal rays; Leptacanthichthys has five
or six (Table 1; fin-ray counts for other oneirodids are given by Pietsch 1974:27-28,
table 23).

Pectoral lobe. — The pectoral lobe (pectoral fin, excluding fin rays) of Puck,
Ctenochirichthys, Leptacanthichthys and, to a lesser degree, Chirophryne (but known
only from small specimens), is unusually long and narrow (Fig. 8; Regan and Tre-
wavas 1932:35-36, fig. 42C, D; Pietsch 1974:29, fig. 55). Unlike that of other onei-
rodids, the first radial (uppermost) is considerably shorter than the second; the car-
tilaginous distal end of the third radial is expanded in an antero-posterior direction to
meet the distal end of the considerably shorter second radial. The fin rays, thus ar-
ticulate along the upper margin of the pectoral lobe (Figs. 9, 11, 14, 16).

A curious division of the distal end of the lowermost radial and its cartilage is
present in the pectoral lobe of both known specimens of Puck (Fig. 8 A), possibly

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No. 297

Figure 6. Lateral view of opercular bones, right side: A. Puck pinnata, SIO H52-363, 66.0
mm; B. Puck pinnata, holotype LACM 34276-1, 46.0 mm; C. Ctenochirichthys longimanus,
paralectotype, BMNH 1932.5.3.20, 36.5 mm; D . Leptacanthichthys gracilispinis, ROM 27284,
54.0 mm; E. Leptacanthichthys gracilispinis, ROM 27293, 43.0 mm; F. Leptacanthichthys gra-
cilispinis, ROM 27274, 41.0 mm; G. Chirophryne xenolophus, SIO 70-306, 22.0 mm.

Figure 7. Pharyngobranchials of Puck pinnata, SIO H52-363, 66.0 mm.

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N. Pacific Ocean Ceratioid Anglerfish

9

Figure 8. Lateral view of pectoral radials, left side: A. Puck pinnata, SIO H52-363, 66.0 mm;
B. Ctenochirichthys longimanus, paralectotype, BMNH 1932.5.3.20, 36.5 mm.

indicating a fusion of two radials (see Pietsch 1972:41-42, fig. 23). All oneirodids
are thought to have three pectoral radials, the possible presence of four radials in Puck
further complicates the already cumbersome diagnosis of the family (Pietsch 1974:30).

Puck, Chirophryne, and Leptacanthichthys have pectoral-fin ray counts ranging
from 18 to 21; Ctenochirichthys has considerably more pectoral- fin rays ranging from
28-30 (Table 1).

Skin Spines. — Skin spines could not be detected microscopically in the single
osteo logical preparation of Puck.

SYSTEMATICS

Key to the “long-pectoraled” genera of the Oneirodidae

IA. Sphenotic and articular spines short, in some cases not piercing skin (Fig. 11);
length of pectoral lobe greater than 15 per cent of SL; pectoral fin rays 27-30 .
Ctenochirichthys Regan and Trewavas 1932.

IB. Sphenotic and articular spines long, piercing skin (Figs. 9, 14, 16); length of

pectoral fin lobe less than 15 per cent of SL; pectoral fin rays 17-21 2.

Table 1

Fin-ray frequencies for “long-pectoraled” genera of Oneirodidae

Dorsal
A 5 61

Anal
4 5 6

Pectoral (both sides )

18 19 20 21 — 28 29 30

Puck pinnatus

2

2

4

Ctenochirichthys longimanus

1 3

3 1

1 2 5

Chirophryne xenolophus

2

2

2 2

Leptacanthichthys gracilispinis

1 7 2

9 1

12 4 7

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No. 297

2A. Length of quadrate spine less than length of mandibular spine (Fig. 5B); dorsal
profile of frontal bones nearly linear (Fig. 16); esca with a single appendage

arising from dorsal surface (Fig. 17); anal fin rays 5-6

Leptacanthichthys Regan and Trewavas 1932.

2B. Length of quadrate spine greater than length of mandibular spine (Fig. 5A);
dorsal profile of frontal bones convex (Figs. 9, 14); esca with more than a
single appendage, either five separate appendages arising from dorsal surface
or three dorsal appendages and a lateral filament (Figs. 10, 15); anal fin rays
4 3.

3A. Length of anterior maxillo mandibular ligament greater than one-half length of
premaxillary, gape of mouth extending beyond eye (Fig. 3C); suboperculum
short and broad, upper end rounded (Fig. 6G); esca without a lateral filament
(Fig. 15) Chirophryne Regan and Trewavas 1932.

3B. Length of anterior maxillo mandibular ligament less than one-half length of
premaxillary, gape of mouth terminating before eye (Fig. 3A); suboperculum
long and narrow, upper end tapering to a point (Fig. 6A, B); esca with a lateral
filament (Fig. 10) Puck new genus.

Puck NEW GENUS

Type species. - Puck pinnata new species

Diagnosis for females. — A member of the Oneirodidae as diagnosed by Pietsch
(1974:30) distinguished by the following combination of characters: snout extremely
short, dorsal profile of frontal bones strongly convex, gape of mouth terminating an-
terior to eye; length of anterior maxillomandibular ligament less than one-half length
of premaxillary (Fig. 3 A); hyomandibular with a double head; anterior end of pter-
ygiophore of illicium exposed, its posterior end concealed under skin; illicium length
10.7 - 18.5 per cent of SL; lower jaw with a small symphysial spine; sphenotic spines
well developed; articular spines present, quadrate spine nearly six times length of
mandibular spine (Fig. 5 A); angular spine absent; vomerine teeth present; pharyn-
gobranchial I absent; pharyngobranchials II and HI present and toothed (Fig. 7); epi-
branchial teeth absent; hypobranchial II present; pectoral-fin lobe long and narrow,
longer than the longest rays of pectoral fin (Figs. 8 A, 9); suboperculum elongate,
slender throughout length, upper end tapering to a point, lower end without anterior
spine or projection (Fig. 6A, B); anal fin with 4 rays; skin naked (embedded dermal
spines cannot be detected microscopically in bleached and stained skin), covering
caudal fin to some distance from fin base.

Puck resembles the oneirodid genera Ctenochirichthys, Chirophryne, and Lep-
tacanthichthys in having an unusually long and narrow pectoral-fin lobe in which the
first radial is considerably shorter than the second, and the second ankylosed to the
distal expansion of the third (Fig. 8 A, B). Puck is clearly distinguished from these
forms, however, by its extremely short snout, strongly convex frontal bones, and
oblique jaw suspension. It is further distinguished from Chirophryne and Leptacan-

1978

N. Pacific Ocean Ceratioid Anglerfish

11

thichthys by having a short anterior maxillomandibular ligament, a short, yet wide
mouth (Figs. 3, 13), and an elongate subopercular bone (Fig. 6); and from Cteno-
chirichthys by its well-developed sphenotic and articular spines (Figs. 9, 1 1, 13).

Males and larvae. — Unknown.

Etymology. — The name Puck, to be treated as a noun of feminine gender, is
taken from Germanic folklore, a minor order of mischievous devils, sprites, goblins
or demons; a devilish trickster.

Puck pinnata new species
Figures 1-3A, 4, 5A, 6A,B, 7, 8A, 9, 10, 13A, 18
Tables 1, 2

Holotype. — LACM 34276-1, 46.0 mm, female; VITYAZ Cruise 19, Station
3199, Sample 123b, western north Pacific, 38° 16' N, 152° 34' E; 6 m diameter
conical ring net fished open with 5350 m of wire out; bottom depth 5420-5350 m;
0230-0545 hr; 16 October 1954.

Paratype. — SIO H52-363, 66.0 mm, female; HORIZO N, between Stations SB
101 and 102; Berner, Juhl and Schaefer, collectors; eastern tropical Pacific, 6° 58'
N, 88° 35' W; 3 m IKMT, fished open between surface and 1464 m; 1 July 1952.

Description of the holotype. — Escal bulb with a stout, rounded, anterodorsally
directed and internally pigmented anterior papilla; a similar, posterodorsally directed
medial papilla without internal pigment; distal ends of anterior and medial papillae
darkly pigmented; an unpigmented, compressed posterior appendage bearing antero-
dorsally a lump of tissue of uncertain morphology; and a tapering, unpigmented lateral
filament on each side, slightly less than length of escal bulb (Fig. 10).

Operculum bifurcate, the two forks forming an acute angle of 25°; length of lower
fork 23.9 per cent of SL, upper fork 72.7 per cent of length of lower fork; length of
suboperculum 40.9 per cent of length of lower fork (Fig. 6A, B).

Teeth as described for family; longest tooth in upper jaw 2.0 mm, in lower jaw
3.0 mm; pelvics absent; caudal fin 9 (2 unbranched - 4 branched - 3 unbranched);
branchiostegal rays 6 (2 + 4).

Figure 9. Holotype of Puck pinnata, LACM 34276-1, 46.0 mm.

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No. 297

Figure 10. Esca of Puck pinnata, holotype, LACM 34276-1, 46.0 mm.

Fin ray counts are given in Table 1, tooth counts and body measurements in
Table 2.

Etymology. — The specific name is derived from the Latin pinnata, meaning
plumed or winged, in reference to the narrow, elongate pectoral fins.

Distribution. — Puck pinnata is known from two metamorphosed females: the
holotype collected from the western north Pacific at 38° 16' N, 152° 34' E, and a
second specimen from the eastern tropical Pacific at 6° 58' N, 88° 35' W (Fig. 18).

Ctenochirichthys Regan and Trewavas

Ctenochirichthys Regan and Trewavas 1932:82, pi. Ill, fig. 3 (type species Ctenochirichthys
longimanus Regan and Trewavas 1932, by original designation).

Diagnosis for Females (based on two metamorphosed females, 12.5 and 36.5
mm). — The genus Ctenochirichthys differs from all other oneirodid genera in having
greatly reduced sphenotic and articular spines that, in some cases do not pierce the
skin, an extremely elongate pectoral-fin lobe greater than 15 per cent of SL, and a

Table 2

Tooth counts and body measurements in percent of SL for “long-pectoraled” genera of Oneirodidae

1978

N. Pacific Ocean Ceratioid Anglerfish

13

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14

Contributions in Science

No. 297

high number of pectoral fin rays, 27 to 30. In addition, Ctenochirichthys is unique
in having the following combination of characters: snout short, dorsal profile of frontal
bones convex; gape of mouth terminating anterior to or beneath eye; length of anterior
maxillomandibular ligament less than one-half length of premaxillary (Fig. 3B); hy-
omandibular with a double head; anterior end of pterygiophore of illicium exposed,
its posterior end concealed under skin; illicium length 24.8 - 31.2 per cent of SL;
lower jaw with a small symphysial spine; angular spine absent; vomerine teqth present;
pharyngo-branchials II and HI present and toothed; epibranchial teeth absent; suboper-
culum elongate, slender throughout length, upper end tapering to a point, lower end
without anterior spine or projection (Fig. 6C); anal fin with 4 or 5 rays; skin presum-
ably naked (bleached and stained skin not available for examination), covering caudal
fin to some distance from fin base.

Diagnosis for males and larvae. — See Bertelsen (1951:95).

Ctenochirichthys longimanus Regan and Trewavas
Figures 3B, 6C, 8B, 11-13B, 18
Tables 1, 2

Dolopichthys heteracanthus Regan 1926:28 (in part).

Ctenochirichthys longimanus Regan and Trewavas 1932:82, pi. Ill, fig. 3 (original description;
two specimens; lectotype, designated by Bertelsen 1951, ZMUC P9297). Bertelsen 1951:75,
94-96, figs. 51-52, table 16 (redescription of type material; males and larvae described;
opercular bones, larval male, adolescent male, and lectotype figured; in key). Grey 1956:251
(synonymy; distribution). Bussing 1965: 222 (misidentification, specimen here referred to
Pentherichthys atratus ). Pietsch 1974:31, 32, 89, table 23 (relationships; in key).
Trematorhynchus multiradiatus Beebe and Crane 1947: 166, text fig. 11 (original description;
single male specimen; CAS-SU 46491). Bertelsen 1951:95, fig. 52B (redescription; com-
parison with all known material; figured).

Material. — Two females, 12.5 - 36.5 mm, two larval males, 4.5 - 5.0 mm, and
a single adolescent male, 11.5 mm.

Figure 11. Paralectotype of Ctenochirichthys longimanus, BMNH 1932.5.3.20, 36.5 mm.

1978

N. Pacific Ocean Ceratioid Anglerfish

15

Lectotype of Ctenochirichthys longimanus. — ZMUC P9297, 12.5 mm; DANA
Station 3548 (2); Gulf of Panama, 7° 06' N, 79° 55' W; 3000 m wire; 1030 hr; 3
September 1928.

Paralectotype of Ctenochirichthys longimanus — BMNH 1932.5.3.20, 36.5 mm;
DANA Station 1206 (5); Gulf of Panama, 6° 40' N, 80° 47' W; 2500 m wire; 1845
hr; 14 January 1922.

Holotype of Trematorhynchus multiradiatus. — CAS-SU 46491, 11.5 mm; East-
ern Pacific ZACA Expedition Station 225, Net No. T-l; 11 miles southwest of Jicaron
Island, Panama, 7° 08' N, 81° 57' W; 1 m diameter conical ring net, 0-910 m; 20
March 1938.

Non-type material. — ZMUC P92795, 5.0 mm; DANA Station 3548(7); Gulf of
Panama, 7° 06' N, 79° 55' W; 100 m wire; 2000 hr; 3 September 1928. ZMUC

Figure 12. Esca of Ctenochirichthys longimanus, lectotype, ZMUC P9297, 12.5 mm.

16

Contributions in Science

No. 297

P92794, 4.5 mm, DANA Station 1141(14); eastern north Atlantic, 34° 15' N, 16° 53'
W; 300 m wire; 2355 hr; 14 October 1921.

Description of females. — Escal bulb with a short, rounded anterior appendage;
a darkly pigmented, raised band of tissue extending over dorso-medial surface and
down onto sides of bulb with a circular unpigmented area on each side; an unpig-
mented, compressed, tapering posterior appendage; lateral filaments absent (Fig. 12).

Operculum bifurcate, the two forks forming an acute angle of 38°; length of lower
fork 21.9 per cent of SL, upper fork 52.5 per cent of length of lower fork; length of
suboperculum 51.2 per cent of length of lower fork of operculum (Fig. 6C).

Teeth as described for family; pelvics absent; caudal fin 9 (2 unbranched-
4 branched - 3 unbranched); branchiostegal rays 6 (2 + 4). Fin ray counts are given
in Table 1, tooth counts and body measurement in Table 2.

Distribution . — Ctenochirichthys longimanus is known from five specimens, four
of which were collected in the Gulf of Panama. The fifth specimen, a larval male,
was taken in the eastern north Atlantic at 34° 15' N, 16° 53' W (Fig. 18).

Chirophryne Regan and Trewavas

Chirophryne Regan and Trewavas 1932:81-82, figs. 131-132 (type species Chirophryne xeno-
lophus Regan and Trewavas 1932, by original designation).

Diagnosis for females (based on two metamorphosed specimens, 11.0 and 22.0
mm). — Chirophryne differs from other oneirodid genera in having the following com-
bination of characters: snout moderate in length, dorsal profile of frontal bones con-
vex; gape of mouth extending past eye; length of anterior maxillomandibular ligament
greater than half length of premaxillary (Fig. 3C); hyomandibular with a double head;
anterior end of pterygiophore of illicium exposed, its posterior end concealed under
skin; illicium length 18.2 - 22.7 per cent of SL; lower jaw with a small symphysial
spine; sphenotic spines well developed; articular spines present, quadrate spine nearly
six times length of mandibular spine; vomerine teeth present; pharyngobranchials II
and III present and toothed; epibranchial teeth present; pectoral-fin lobe long and nar-
row, longer than the longest rays of pectoral fin; suboperculum short and broad, upper
end rounded, lower end without anterior spine or projection (Fig. 6G); anal fin with
4 rays; skin presumably naked (bleached and stained skin not available for exami-
nation), covering caudal fin to some distance from fin base.

Males and larvae. — Unknown.

Chirophryne xenolophus Regan and Trewavas
Figures 3C, 6D, 13C, 14, 15, 18
Tables 1, 2

Chirophryne xenolophus Regan and Trewavas 1932:81-82, figs. 131-132 (original description;
single specimen; holotype ZMUC P9296). Bertelsen 1951:75, 94, fig. 50 (redescription of
holotype; opercular bone figured; in key). Grey 1956:250-251 (synonymy; distribution).
Pietsch 1974:31, 32, 89, table 23 (relationships; in key).

1978

N. Pacific Ocean Ceratioid Anglerfish

17

Material. — Two adolescent females, 11.0 - 22.0 mm.

Holotype. — ZMUC P9296, 11.0 mm, DAN A Station 3731(12); South China Sea,
14° 37' N, 119° 52' E; 2500 m wire; 0200 hr; 17 June 1929.

Non-type material. — SIO 70-306, 22.0 mm; ANTIPODES Expedition Station
4-51A, Trawl 1; 32° 10' N, 136° 05' E; 0-1400 m; 2355-0746 hr; 28-29 August 1970.

Description of 22.0 mm female. — Escal bulb with an unpigmented, tapering an-
terior appendage connected by a thin membrane to an internally-pigmented antero-

C D

Figure 13. A. Puck pinnata, holotype, LACM 34276-1, 46.0 mm; B. Ctenochirichthys lon-
gimanus, paralectotype, BMNH 1932.5.3.20, 36.5 mm; C. Chirophryne xenolophus, SIO 70-
306, 22.0 mm; D. Leptacanthichthys gracilispinis, LACM 33625-2, 56.0 mm.

18

Contributions in Science

No. 297

Figure 14. Chirophryne xenolophus, SIO 70-306, 22.0 mm.

Figure 15. Esca of Chirophryne xenolophus, SIO 70-306, 22.0 mm.

dorsal appendage that is darkly pigmented on distal tip except for a small, circular,
unpigmented area on each side; a pair of bilaterally placed, unpigmented medial ap-
pendages; an unpigmented, compressed, posterior appendage bearing distally a rounded
lump of tissue that tapers to a point; lateral filaments absent (Fig. 15; Regan and
Trewavas 1932:82 fig. 132).

1978

N. Pacific Ocean Ceratioid Anglerfish

19

Operculum bifurcate, the two forks forming an acute angle of 37°; length of lower
fork 25.4 per cent of SL, upper fork 55.2 per cent of length of lower fork; length of
suboperculum 32.5 per cent of length of lower fork of operculum (Fig. 6G).

Teeth as described for family; longest tooth in upper jaw 0.5 mm; in lower jaw
1.0 mm; pelvics absent; caudal fin 9 (2 unbranched - 4 branched - 3 unbranched);
branchiostegal rays 6 (2 + 4).

Fin ray counts are given in Table 1, tooth counts and body measurements in
Table 2.

Distribution. — Chirophryne xenolophus is known from two adolescent females:
the holotype collected from the South China Sea and a second specimen from off
Japan at 32° 10' N, 136° 05' E (Fig. 18).

Leptacanthichthys Regan and Trewavas

Dolopichthys (subgenus Leptacanthichthys) Regan and Trewavas 1932:66, 80, fig. 128 (genus
Dolopichthys broadened to incorporate five subgenera, amended by Bertelsen 1951; type
species Dolopichthys gracilispinis Regan 1925, by monotypy).

Leptacanthichthys Bertelsen 1951:74, 94, fig. 49 (subgenus Leptacanthichthys given generic
status; type species Dolopichthys gracilispinis Regan 1925, by monotypy).

Diagnosis for females (based largely on osteological evidence presented by
Pietsch 1974). — The genus Leptacanthichthys differs from other one irodids by having
a well-developed mandibular spine that is considerably longer than the quadrate spine
(Fig. 5B). In addition, Leptacanthichthys is unique in having the following combi-
nation of characters: snout long, dorsal profile of frontal bones nearly linear; gape of
mouth extending past eye; length of maxillomandibular ligament greater than half
length of premaxillary (Fig. 3D) hyomandibular with a double head; anterior end of
ptergiophore of illicium exposed, its posterior end concealed under skin; illicium
length 19.2-24.1 per cent of SL; lower jaw with a small symphysial spine; sphenotic
spines well developed; angular spines absent; vomerine teeth present; pharyngobran-
chial I absent; pharyngobranchials II and III present and toothed; epibranchial teeth
absent; hypobranchial II present; pectoral-fin lobe long and narrow, longer than the
longest rays of pectoral fin; suboperculum short and broad, upper end rounded to
bluntly pointed, lower end without anterior spine or projection (Fig. 6D-F) anal fin
with 5 rays, rarely 6; skin naked (embedded dermal spines cannot be detected mi-
croscopically in bleached and stained skin), covering caudal fin to some distance from
fin base.

Diagnosis for males (based on a single known male in late metamorphosis, par-
asitically attached to a sexually mature female, LACM 33625-2; Pietsch 1976 figs.
2-5). — Posterior nostril well separated from eye; upper end of suboperculum rounded;
6 lower denticles; gill cover pigmented with slightly darker pigmentation along pos-
terior margin of suboperculum; dorsal pigment restricted to upper part of body ex-
tending beneath base of dorsal fin and just past anterior base of anal fin with a more
heavily pigmented dorsal and ventral group of melanophores near hypural plate; per-
itoneum pigmented.

Larvae. — Unknown.

20

Contributions in Science

No. 297

Leptacanthichthys gracilispinis (Regan)

Figures 3D, 5B, 6E-H, 13D, 16-18
Tables 1 , 2

Dolopichthys gracilispinis Regan 1925:563 (original description; two specimens; lectotype des-
ignated by Bertelsen 1951, ZMUC P9295). Regan 1926:27, 30, pi. V, fig. 2 (brief de-
scription after Regan 1925; in key).

Dolopichthys (Leptacanthichthys) gracilispinis Regan and Trewavas 1932:66, 80 fig. 128 (genus
Dolopichthys broadened to incorporate five subgenera, amended by Bertelsen 1951; de-
scription after Regan 1925, 1926; in key).

Leptacanthichthys gracilispinis Bertelsen 1951:74, 94, fig. 49 (subgenus Leptacanthichthys
given generic status; description; in key). Grey 1956:250 (synonymy; distribution). Pietsch
1974:16-32, 82, 86-89, Figs. 22, 34, 39c, 42, 51c, 55, 59, 103, 104, tables 23, 24 (os-
teology; relationships, esca figured; in key).

Material. — Thirteen females, 10.5-56.0 mm, and a parasitic male in late met-
amorphosis, 7.5 mm.

Lectotype. — ZMUC P9295, 52.0 mm; DANA Station 1206(3); Gulf of Panama,
6° 40' N, 80° 47' W; 3500 m wire; 1845 hr; 14 January 1922.

Paralectotype. — BMNH 1925.8.11.14, 43.0 mm, DANA Station 1358(5); north
Atlantic, 28° 15' N, 56° 00' W; 3000 m wire; 1530 hr; 2 June 1922.

Non-type material. — ROM 27284, 54.0 mm \ BRANDAL Tow 20; 43° 23' N, 52°
30' W; Engel trawl, 0-1050 m; 25 July 1968. ROM 27293, 43.0 mm; BRANDAL
Tow 22; 44° 00' N, 57° 52' W; Engel trawl, 0-1000 m; 26 July 1968. ROM 27274,
41.0 mm; BRANDAL Tow 14; 46° 00' N, 44° 30' W; Engel trawl, 0-1000 m; 18 July
1968.

LACM 33625-2, female, 56.0 mm, with parasitic male, 7.5 mm; VELERO IV
Station 19009; 8° 10' N, 86° 00' W; 3 m IKMT, 0-750 m; 0625 hr; 24 May 1973.
LACM 34275-1, 27.0 mm; VITYAZ Cruise 24, Station 3573, Sample 50; 38° 04' N,
144° 13' E; 1.6 m conical ring net, 0-3000 m; bottom depth 5660-5760 m; 0950-1530
hr; 4 May 1957. LACM 32776-1, 22.0 mm; TERITU Cruise Blood, Sweat, and Tears,
Sample 70-7-6; 21° 20' -30' N, 158° 20-30' W; 2 m IKMT, 0-1250 m; 0746-1200 hr;
6 July 1970.

Figure 16. Leptacanthichthys gracilispinis, LACM 33625-2, 56.0 mm. Stalk of tissue protrud-
ing from belly bears embedded bones of upper jaw of a parasitic male (see Pietsch 1976).

1978

N. Pacific Ocean Ceratioid Anglerfish

21

Figure 17. Esca of Leptacanthichthys gracilispinis, LACM 32776-1, 22.0 mm.

SIO 60-235, 13.0 mm; TETHYS Expedition; 0° 08.5' S, 138° 50.9' W; 0-2100
m; 3 - 4 July 1960. SIO 68-476, 10.5 mm; STYX Expedition VII-40; 22° 00' N,
171° 40' E; 3 m IKMT, 0-1250 m; bottom depth 3660 m; 0640-1300 hr; 15 September

1968.

CAS-SU 43423, 14.0 mm; GLADISFEN Net 874; 32° 12' N, 64° 36' W; 11
September 1930.

IOS uncatalogued, 12.5 mm; DISCOVERY H 8281-37; 31° 48' N, 63° 37' W;
Rectangular Midwater Trawl with closing device, 1240-1265 m; 0200-0500 hr; 19
March 1973.

MCZ 50705, 12.5 mm; ATLANTIS II Cruise 49, RHB 1939; 40° 22' N, 58° 51'
W; 3 m IKMT, 0-1040 m; 1235-1440 hr; 8 July 1969.

Description of females. — Escal bulb with a darkly pigmented streak on dorsal
surface, and an unpigmented, compressed posterior appendage; anterior and lateral
appendages absent (Fig. 17; Regan and Trewavas 1932:80 fig. 128).

22

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No. 297

Figure 18. Distributions of “long-pectoraled” oneirodid genera. A single symbol may indicate more than one capture.

1978

N. Pacific Ocean Ceratioid Anglerfish

23

Operculum bifurcate, the two forks forming an acute angle of 18° to 27°; length
of lower fork 18.1-19.5 per cent of SL; length of lower fork 64.4-70.5 per cent of
length of lower fork; length of suboperculum 29.5-37.5 per cent of length of lower
fork of operculum (Fig. 6D-F).

Teeth as described for family; D. 4-6, of 10 specimens counted one had D.4;
A. 5-6, P. 18-21, of 8 specimens counted, one had P.18; pelvics absent; caudal fin
9 (2 unbranched - 4 branched - 3 unbranched); branchiostegal rays 6 (2 + 4).

Fin ray counts are given in Table 1, tooth counts and body measurements in
Table 2.

Distribution. — In the Atlantic Ocean, L. gracilispinis seems to be restricted to
waters off the continental slope of North America where about one-half of the known
material has been collected between 28° and 46° N, as far east as 44° W. The re-
maining material is from the north Pacific where it ranges from off Japan to the Gulf
of Panama (Fig. 18). Ninety-two per cent of the known material was captured by gear
fished below 1000 m; 62 per cent by gear fished below 1200 m.

DISCUSSION

The ceratioid family Oneirodidae is by far the largest and most diverse of the 1 1
deepsea anglerfish families, but probably remains the least well known. With the
addition of Puck, 15 oneirodid genera are now recognized. Eleven of these genera
consist of a single species and of these, nine are based on only one or two adolescent
or adult females. The addition of yet another monotypic genus based on two female
specimens is justified, however, as this new form clearly cannot reasonably be placed
within any previously described genus. Puck , Ctenochirichthys, Chirophryne, and
Leptacanthichthys share an unusually elongate pectoral- fin lobe that bears the fin rays
along its dorsal margin. A similar arrangement is not found in any other ceratioid; it
is mainly because of the common occurrence of this derived character complex that
these four genera are thought to form a natural assemblage. Within this group of four
genera, Puck appears to be most closely related phylogenetically to Ctenochirichthys .
These two genera share a modification of the jaw mechanism in which the gape of
the mouth is considerably shorter, but at the same time, wider, relative to other onei-
rodids. In addition, Puck and Ctenochirichthys share an elongate suboperculum, un-
like the short and broad suboperculum of Chirophryne and Leptacanthichthys .

Puck appears to be intermediate in many ways between more generalized onei-
rodid genera, especially Danaphryne, and the other three, more specialized “long-
pectoraled” genera. Puck is similar to Danaphryne and unlike Leptacanthichthys in
the shape and relative size of the bones of the ethmoid region, in the shape and position
of the frontals, and in the length and depth of the cranium and elements of the man-
dibular, palatine, and hyoid arches. Puck, on the other hand, shares the specialized
pectoral lobe with Ctenochirichthys, Chirophryne, and Leptacanthichthys as well as
the modification of the jaw mechanism of Ctenochirichthys.

Although apparently most closely related to Puck, probably the least derived of
the four “long-pectoraled” genera, Ctenochirichthys is considered the most derived

24

Contributions in Science

No. 297

member of the group in having reduced sphenotic and articular spines, a reduced
number of vomerine teeth, a considerably more elongate pectoral lobe, a secondarily
derived increase in the number of pectoral-fin rays, and a more elongate illicium.

Little can be said concerning relationships of Chirophryne, known from only two
small, adolescent females. It is perhaps intermediate between Puck and Ctenochirich-
thys on one hand, with which it shares convex frontal bones, and Leptacanthichthys
on the other, with which it shares a similar jaw mechanism and similarly- shaped sub-
opercular bones.

Leptacanthichthys is derived and unique among “long-pectoraled” oneirodid
genera in having a nearly linear profile of the frontal bones; it is unique among onei-
rodids in having a well-developed mandibular spine that is considerably longer than
the quadrate spine.

ACKNOWLEDGMENTS

I thank E. Bertelsen, Robert J. Lavenberg and Camm C. Swift for critically
reading the manuscript. The following people and institutions provided study material:
E. Bertelsen (ZMUC); Robert J. Lavenberg and Jerry W. Neumann (LACM); Richard
H. Rosenblatt (SIO); Alwyne Wheeler (BMNH); Nigel Merrett and Julian Badcock
(IOS); William N. Eschmeyer and Tomio Iwamoto (CAS); Thomas A. Clarke (Hawaii
Institute of Marine Science); W. B. Scott (ROM); and Karsten E. Hartel (MCZ).
Special thanks go to N. V. Parin, of the Institute of Oceanography of the Academy
of Sciences of the USSR, Moscow, for his gift of the holotype of Puck pinnata to the
Natural History Museum of Los Angeles County; and to Richard Rosenblatt for al-
lowing me to clear and stain the second known specimen of Puck. Figures 9-11, 13-
14, and 16 were made by Patricia Chaudhuri. The work was supported by a grant
from the National Science Foundation, No. GB-40700. Partial assistance from the
William F. Milton Fund of Harvard University and the Johannes Schmidt Fund of the
University of Copenhagen is also gratefully acknowledged.

LITERATURE CITED

Beebe, W. and J. Crane 1947. Eastern Pacific Expedition of the New York Zoological So-
ciety. XXXVII. Deep-sea ceratioid fishes. Zoologica, N.Y. 31(1 1): 1 5 1 — 1 82.

Bertelsen, E. 1951. The ceratioid fishes. Ontogeny, taxonomy, distribution and biology. Dana
Rept. 39:1-276.

Bradbury, M. G. 1967. The genera of batfishes (family Ogcocephalidae). Copeia 1967:
399-422.

Bussing, W. A. 1965. Studies of the midwater fishes of the Peru-Chile Trench. Biol. Antarctic
Seas II. Amer. Geophys. Union Res: Ser. 5:185-227.

Garman, S. 1899. Reports on an exploration of the west coasts of Mexico, Central and South
America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish
Commission steamer “Albatross” during 1891, Lieut. Commander Z. L. Tanner, U.S.N.,
commanding. XXVI. The fishes. Mus. Comp. Zool., Mem. 24:1-431.

Grey, M. 1956. The distribution of fishes found below a depth of 2000 meters. Fieldiana, Zool.
36(2):75-337.

1978

N. Pacific Ocean Ceratioid Anglerfish

25

Pietsch, T. W. 1972. A review of the monotypic deep-sea anglerfish family Centrophrynidae:
taxonomy, distribution and osteology. Copeia 1972:17-47.

1974. Osteology and relationships of ceratioid anglerfishes of the family Oneirodidae

with a review of the genus Oneirodes Liitken. Nat. Hist. Mus. Los Angeles Co., Sci. Bull.
18:1-113.

1976. Dimorphism, parasitism, and sex: reproductive strategies among deep-sea cer-
atioid anglerfishes. Copeia 1976: 781-793.

Regan, C. T. 1925. New ceratioid fishes from the N. Atlantic, the Caribbean Sea, and the Gulf
of Panama, collected by the “Dana.” Ann. Mag. Nat. Hist., Ser. 8, 8(62):561-567.
. 1926. The pediculate fishes of the suborder Ceratioidea. Dana Oceanog. Rept.

2:1-45.

Regan, C. T. and E. Trewavas. 1932. Deep-sea anglerfish (Ceratioidea). Dana Rept. 2:1-
113.

Taylor, W. R. 1967. An enzyme method of clearing and staining small vertebrates. U.S. Nat.
Mus., Proc. 122:1-17.

Accepted for publication January 16, 1977.

507 s 13
C%L%G%

NUMBER 298
MAY 22, 1978

REPRODUCTION IN MACROGEN IOGLOTTUS ALIPIOI CARVALHO

(ANURA, LEPTODACTYLIDAE)

By J. Paul Abravaya and James F. Jackson

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCICNCC

Published by the NATURAL HISTORY MUSEUM
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SERIAL PUBLICATIONS OP THE
NATURAL HISTORY MUSEUM OP LOS ANGELES COUNTY

Prior to November 30, 1973, publications of the Natural History Museum have appeared
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should be sent to the Editor, Museum Publications, Natural History Museum of Los Angeles
County, 900 Exposition Boulevard, Los Angeles, California 90007.

Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

REPRODUCTION IN MACROGEN IOGLOTTUS ALIPIOI CARVALHO
(ANURA, LEPTODACTYLIDAE)1

By J. Paul Abravaya2 and James F. Jackson3

Abstract: Mating behavior, mating call, and larval morphology are described
for the leptodactylid fro g Macrogenioglottus alipioi Carvalho. The mating behavior
is conducted in a way that would be advantageous in a species in which the number
of eggs oviposited per female is great but the number of ovipositing females per pond
is small. Feeding behavior is modified for the capture of slow moving prey such as
snails and earthworms. The sonogram of the mating call of M. alipioi is similar to
that of Odontophrynus americanus. The tadpoles of Macrogenioglottus are similar
to those of Odontophrynus , having the same tooth row formula and similar arrange-
ment of labial papillae. They differ, however, in the position of the spiracle and the
vent. A review of the taxonomic history of M. alipioi combined with new data sug-
gests a close relationship between Macrogenioglottus and Odontophrynus.

INTRODUCTION

Although Macrogenioglottus alipioi Carvalho was described 30 years ago (Car-
valho 1946) and has been considered of no small interest to anuran phylogeny (Reig
1972; Duellman 1975), little on its biology has been published. We report our ob-
servations on reproduction of the species made in 1974 at the Reserva Biologica Nova
Lombardia, Municipio de Santa Teresa, Espirito Santo, Brazil. The vegetation of the
area was classified as Subtropical Lower Montane Moist Forest by the Holdridge
(1967) system. Because of the extremely dissected topography, the only lentic water
(other than that in tank bromeliads) is found in oxbow ponds along the valley streams.
These ponds are empty much of the year and receive significant amounts of water only
when their progenitory streams overflow after heavy rains characteristic of the wet
season. Choruses of Macrogenioglottus alipioi were heard only after extremely heavy
rains which resulted in such flooding.

Our observations were made at an oxbow pond 30 by 6 m and less than a meter
deep. The water was stained by tannic acids and on the pond bottom was a deep layer
of organic mud and decaying plant material. The elevation of the pond is 810 m.

The pond was visited several times a month between June 1973 and November
1974. No reproductive activity was noted on rainless days or nights, but activity was

Review Committee for this Contribution
Robert L. Bezy
W. Ronald Heyer
Roy W. Me Diarmid
John W. Wright

2Research Associate in Mammalogy, Natural History Museum of Los Angeles County, and De-
partment of Biology, California State University, Northridge, California 91234.

department of Biology, University of Southwestern Louisiana, Lafayette, Louisiana 70504.

2

Contributions in Science

No. 298

noted on the two visits we made during nights of substantial rain, and during one day
after a heavy rain.

Specimens of adult Macro genioglottus alipioi were deposited in the Museu de
Biologia “Prof. Mello Leitao,’’ Santa Teresa, Brazil. Eighteen tadpoles were depos-
ited in the Natural History Museum of Los Angeles County (LACM 121739).

REPRODUCTIVE BEHAVIOR

On the night of 4 January 1974 a chorus of 6 male M. alipioi was heard. All
males were stationed on the bank within a meter of the water. No females were found
at the pond, but a female was watched for an hour as she walked several hundred
meters along a dirt road toward the chorus. When she reached the point on the road
nearest the pond, she turned uphill and proceeded in the opposite direction from the
pond.

On 10 February 1974 observations were made at the pond from 19:30 until 01:30
(local time) of the next day. Three calling males were stationed on the bank at the
water’s edge. One male once moved into the pond and called while standing in water
2-3 cm deep. An amplexed pair was located in the pond at 20:30. Amplexus is ax-
illary, the male’s forelegs clasped around the female one-third to one-fourth the dis-
tance between her fore- and hindlegs, and his forefeet placed in her axillae. Initially
the pair floated quietly and moved only every five minutes; later they moved once or
more each minute. The female initiated changes of location by swimming or walking
one to several meters dragging the male behind. When they reached a new site, ovi-
position usually occurred. When she stopped, the male flexed his forelegs and pulled
himself forward on top of the female. While in this position, he flexed his hindlegs
and brought his hindfeet together at the female’s vent. The hindfeet were then pushed
directly backward in a single stroke that started slowly and then accelerated, as if the
male were conducting eggs or sperm posteriorly. During or after this movement, the
male slid backwards off the female. Then the male kicked several times rapidly; these
kicks were sometimes between the female’s hindlegs but usually lateral to them.
Whether the male is distributing sperm over the eggs or whether he is distributing the
eggs through the water, or both, was impossible to determine. The number of eggs
oviposited at each site was not determined but probably is small, perhaps less than
20. Eggs were laid in short strands and small clusters, 2-eggs thick. They adhered to
vegetable debris and did not float. The amplexed pair was collected at 23:30 and laid
fertilized eggs in the plastic collecting bag. A second female that presumably had not
oviposited was found walking toward the pond. She was collected and found to contain
approximately 3650 eggs.

Although we have no data on tadpole vagility or survivorship, for heuristic pur-
poses we hypothesize three possible adaptive functions of the multiple oviposition.
The frequent changes of oviposition sites and the deposition of eggs in small batches
could function to reduce the probability that all the eggs would be lost due to pre-
dation or desiccation. Spreading the tadpoles around the pond could possibly also
function to minimize competition between siblings. Such a function would be advan-
tageous in a species where the number of eggs oviposited per female is great but the

1978

Reproduction in Macrogenioglottus Alipioi Carvalho

3

number of ovipositing females per pond is small, as it appears to be inM, alipioi judg-
ing from our observations and the fact that the type specimens were a single pair col-
lected in amplexus (Carvalho 1946). In a species with numerous amplexing pairs per
pond, there would be little advantage to such egg dispersal because tadpoles produced
by other pairs would be hatching throughout the pond.

MATING CALL

The call of M. alipioi sounds most like a fog horn. Figure 1 illustrates the son-
ogram of calls recorded on a Uher model 4000 at an ambient temperature of 17°-20°C
on 10 February. The call is a series of pulsed notes, each note lasting 0.24-0.27
seconds. The number of notes in the call varies from one to several dozen. When
repeated, the notes are separated by intervals of 0.54-0.58 seconds. Maximum sound
energy of the moderately well-tuned note is spread over the frequency range 230-800
hz. In the frequency range 800-11 80 hz exists a component of weaker intensity with a
pulse rate of 195-230 pulses per second. The beginning and end of the note on the
sonogram are slightly less dark than the middle, indicating weak intensity modulation.
The low frequency-low intensity “echo” that appears between notes on the sonogram
(Fig. 1) probably originated from another individual in the chorus. The males alternate
their calls to form duets and triplets.

NATURAL HISTORY

Fertilized eggs with their gelatinous envelopes averaged 2.2 mm in diameter.
The tadpoles hatched 48-70 hours after fertilization. In the laboratory, tadpoles grew
even when the only nutrient provided was organic muck from a pond bottom. They
refused to eat meat but readily accepted fruit of avocado and papaya. The tadpoles
stayed near the bottom of the aquarium and did not swim much. This inactivity may
serve a protective function, since the pond had a high density of predaceous Cera-
tophrys tadpoles. Three tadpoles transformed at ages of 86, 95, and 127 days. This
rate of development is consistent with apparent ecological necessities, since the oxbow
ponds are nearly dry by May. The snout- vent lengths of the transformed frogs were
19.9, 19.8, and 21.0 mm.

Stomachs of 5 adult M. alipioi were examined. Two were empty; one contained
2 earthworms; one contained a small snail shell; and one contained the central spiral
of a snail shell and part of an earthworm. An adult maintained in the laboratory ate
snails and earthworms, appearing to prefer the former. It attempted to eat roaches but
was not often successful in picking them up. When a M. alipioi recognized a potential
food item, presumably by its movement, it would walk close, open its mouth, and
pick up the item through a slow extrusion of the tongue. Neither the very deliberate
feeding movements nor the dentition and lingual morphology of M. alipioi appear
adapted for capturing fast-moving prey. Consequently, we believe that our sample of
stomach contents, although limited, accurately reflects the dietary habits of the
species.

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Contributions in Science

No. 298

Figure 1. Sonogram of mating call of Macrogenioglottus alipioi Carvalho. 10 February 1974; Santa Teresa, Espirito, Santo, Brazil; ambient temp-
erature 17-20 C.

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Reproduction in Macrogen ioglottus Alipioi Carvalho

5

DESCRIPTION OF TADPOLE

The tadpole of M. alipioi is illustrated in figure 2. The tadpole is characterized
as follows (terminology of Altig 1970): sinistral spiracle; dextral anus; complex
mouthparts; labial papillae well developed laterally and complete along posterior la-
bium, absent on median portion of anterior labium; labia strongly emarginate laterally;
tooth rows %; second anterior tooth row with a broad A-2 gap; first posterior tooth
row with narrow P-1 gap; denticles fine and short; upper jaw narrow and smooth;
lower jaw narrow and toothed laterally; eyes and nostrils dorsal, eyes directed dorsally;
body elliptical in dorsal view; somewhat depressed dorso-ventrally, greatest depth .35-
.43 of standard length; dorsal and lateral surface of body brownish-grey, reticulated
with non-pigmented areas; venter transparent; tail musculature pigmented with a series
of dark grey spots or bars along dorsal portion and with irregular dark grey reticulation
on ventral portion; margins of tail fin spotted irregularly with dark grey; standard
length 12-18 mm; tail relatively short, 1.24-1.40 of standard length. The tadpole in
Figure 2 is at stage 30 (Gosner 1960); standard length 13.9 mm, tail 21.5 mm, body
width 9.8 mm. Tadpoles at stage 25 show considerable variation in growth; four spec-
imens examined ranged from 4.5 mm to 11.5 mm in standard length.

EVOLUTIONARY RELATIONSHIPS

In the description of Macro genioglottus, Carvalho (1946) assigned the genus to
the Ceratophryidae (=Ceratophrydidae) and considered Odontophrynus its closest rel-
ative. Since then the relationship of Macro genioglottus to Odontophrynus has been
a subject of considerable disagreement. Lynch (1971) synonymized Macro genioglot-
tus with Odontophrynus in the leptodactylid subfamily Telmatobiinae which was sep-
arated from the subfamily Ceratophryinae. From the appendix of his paper one would
assume that this synonymy was made without examination of specimens of Macro-
genioglottus . This presumably is the reason that some of the characters in his diag-
nostic definition of Odontophrynus (sensu lato) actually exclude Macro genioglottus .
For example, Macro genioglottus is considerably larger (Carvalho 1946) and falls out-
side Lynch’s size range for Odontophrynus , and the inner metatarsal tubercle of Mac-
ro genioglottus, though enlarged, is not spade-like. Reig (1972) on the other hand
made a detailed anatomical comparison of Macro genio glottus , Odontophrynus, Cer-
atophrys, and Bufo. He concluded that similarities between Macro genio glottus and
Odontophrynus are convergent and that Macro genioglottus resembles a hypothetical
taxon representing the ancestral bufonoid stock. He proposed elevating Macro gen-
io glottus to familial rank placing it together with the Bufonidae, Atelopodidae and
Ceratophrynidae in the superfamily Bufonoidea, retaining Odontophrynus in the
Leptodactyloidea.

Martin (1972) used evidence from laryngeal anatomy and function, and from
karyotypes (Bogart 1967) to suggest that Odontophrynus, as well as Macrogenio-
glottus, are close to the base of the bufonid -Atelopus radiation. He felt that anatomical
evidence (Reig 1972) actually supported this conclusion and that Reig’s suggestion
of convergence between Macro genio glottus and Odontophrynus was untenable.

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Contributions in Science

No. 298

Figure 2. Stage 30 larva of Macro genioglottus alipioi Carvalho (LACM 121739).

1978

Reproduction in Macrogen ioglottus Alipioi Carvalho

7

Heyer (1975) did not examine specimens of Macro genioglottus but using the data
provided by Reig (1972) suggested that within the New World Leptodactylidae Mac-
ro genioglottus has affinities with Ceratophrys, Lepidobatrachus , Odontophrynus
and Proceratophrys. He did not consider Macro genioglottus and Odontophrynus to
be congeneric.

The tadpole of Macro genioglottus is quite similar to those of Odontophrynus
examined by us and described by Savage and Cei (1965). It has the same tooth row
formula, a similar arrangement of labial papillae and the labia are emarginate as in
O. occidentalis and O. americanus.

It differs principally by having a shorter tail (1.24-1.40 standard length in Mac-
ro genioglottus versus 1.30-2.15 in Odontophrynus ) and finer denticles. Viewed lat-
erally, the spiracle opening of Macro genioglottus is clearly at a level dorsal to the
mouth, whereas in Odontophrynus the spiracle opening is at the same level as the
mouth. There appears to be confusion regarding the position of the vent in Odonto-
phrynus. Savage and Cei (1965) stated that it is median. Lynch (1971:26) called the
vent dextral for the genus Odontophrynus but gave a median vent as a character diag-
nostic for the tribe Odontophrynini (Lynch 1971:131). After examining tadpoles of
Odontophrynus americanus (LACM 28060), O. cultripes (LACM 28059), and O.
occidentalis (LACM 28068), we consider all to be weakly dextral. Regardless of
whether the vent of Odontophrynus is considered median or dextral, it is clearly less
extremely dextral than the vent of Macro genioglottus .

The mating calls of Odontophrynus americanus and O. occidentalis have been
described by Barrio (1964). These calls have similarities to that of Macro genioglottus
in consisting of several unmodulated notes repeated at regular intervals and in having
a dominant frequency between 400-1200 hz. The call ofO. occidentalis is least similar
since it has a very short interval between the notes (about .04 seconds) and is thus
a trill. The inter-note interval of O. americanus (about 0.4 seconds) is closer to that
of Macrogenioglottus . Both Odontophrynus species differ from Macrogenioglottus in
producing calls with complex harmonics. Of the two, O. americanus is again the
closer to Macrogenioglottus by having a less accentuated harmonic structure, sug-
gesting that the pulse rate is similar to that of Macrogenioglottus .

In summary, we feel that the weight of taxonomic evidence indicates a close
relationship between Macrogenioglottus and Odontophrynus and that new evidence
from larval morphology and mating call support this conclusion. However, to include
Macrogenioglottus in the genus Odontophrynus, as presently known, would seem to
violate the homogeneity traditionally expected of a genus of four or five species.

ACKNOWLEDGMENTS

We are grateful to Dr. Jacques Vielliard, Laboratoire de Zoologie, Ecole Nor-
male Superieure, Paris, for recording the mating call and furnishing the sonogram.
We thank Andrew Starrett, Anthony Gaudin, W. Ron Heyer and John Wright for
useful criticisms of the paper. To Robert Bezy we owe special thanks for his critiques

Contributions in Science

No. 298

and help in preparing the manuscript. Field work was supported by a grant from
Instituto Brasileiro de Desenvolvimento Florestal to Museu de Biologia Mello-Leitao,
Santa Teresa, Espirito Santo. We thank Augusto Ruschi for aid during our stay at
Museu Mello-Leitao, Mary Butler for preparing drawings of tadpoles, Marianne
Hata and Terri Togiai for typing the manuscript.

RESUMO

O comportamento nupcial, o grito nupcial e a morfologia larval sao descritos
para o anuro Macrogenioglottus alipioi Carvalho (Leptodactylidae). Na Reserva Nova
Lombardia, Santa Teresa, Espirito Santo, Brazil, o cruzamento ocorre em pequenas
lagoas formadas pelo transbordamento dos corregos durante a esta^ao chuvosa. O
numero de pares que cruzam em cada lagoinha e baixo. Amplexus e axilario. Cada
femea poe mais do que tres mil ovos, mas esses sao distribuidos pela lagoinha em
pequenas por^oes em lugares distintos. O grito nupcial e baixo (frequencia 230-800
hz tern for^a sonora maxima) e e repetido varias vezes. A duragao do grito e de .24-
.27 segundos e o intervalo entre os gritos e de .54-. 58 segundos. O girino e semelhante
ao girino de Odontophrynus , mas existem diferengas na localizafao do espiraculo e
do anus. Observagoes no laboratorio e o exame de conteudos estomacais indicam que
os adultos alimentam-se de caracois e minhocas. Tanto o grito nupcial como a mor-
fologia do girino de Macrogenioglottus sao semelhantes aos de Odontophrynus, mas
diferen^as na morfologia adulta exigem que se os mantenham em generos separados.

LITERATURE CITED

Altig, R. 1970. A key to the tadpoles of the continental United States and Canada. Herpeto-
logica, 26:180-207.

Barrio, A. 1964. Characteres eto-ecologicos diverenciales entre Odontophrynus americanus
(Dumeril et Bibron) y O. occidentalis (Berg) (Anura, Leptodactylidae). Physis, 24:385—
390.

Bogart, J. P. 1967. Chromosomes of the South American amphibian family Ceratophridae with
a reconsideration of the taxonomic status of Odontophrynus americanus . Canadian Jour.
Genet. Cytol., 9:531-542.

Carvalho, A. L. de. 1946. Um novo genero de Ceratofridideo do sudeste baiano. Bol. Museu
Nac., N. S. Zool., Rio de Janeiro, 73:1-5.

Duellman, W. E. 1975. On the classification of frogs. Occas. Papers Mus. Nat. Hist. Univ.
Kansas, 42:1-14.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on
identification. Herpetologica, 16:183-190.

Heyer, W. R. 1975. A preliminary analysis of the intergeneric relationships of the frog family
Leptodactylidae. Smithsonian Contr. Zool., 199:1-55.

Holdridge, L. R. 1967. Life zone ecology. Tropical Science Center, San Jose, Costa Rica.

Lynch, J.D. 1971. Evolutionary relationships, osteology, and zoogeography of leptodactylid
frogs. Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 53:1-238.

1978

Reproduction in Macrogenioglottus Alipioi Carvalho

9

Martin, W. F. 1972. Evolution of vocalization in the genus Bufo\ pp. 279-309 in Evolution
in the genus Bufo (W. F. Blair, ed.), Univ. of Texas Press, Austin and London, 460 pp.
Reig, O. A. 1972. Macrogenioglottus and the South American bufonid toads; pp. 14-36 in
Evolution in the genus Bufo (W. F. Blair, ed.), Univ. of Texas Press, Austin and London,
460 pp.

Savage, J. M. and J. M. Cei. 1965. A review of the leptodactylid frog genus, Odontophrynus .
Herpetologica, 21:178-195.

Accepted for publication February 18, 1977.

50%W

NUMBER 299
JUNE 16, 1978

?

I

Nl|

A NEOGENE SECTION
NORTHEASTERN SAN CLEMENTE ISLAND, CALIFORNIA

By Takeo Susuki and Carol J. Stadum

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTRIBUTIONS IN SCIENCE

Published by the NATURAL HISTORY MUSEUM
OF LOS ANGELES COUNTY
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SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

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CONTRIBUTIONS IN SCIENCE contain articles in the earth and life sciences, presenting
results of original research. Emphasis is intended principally for papers allied to biosystematic
research, but other subjects and review-oriented ones will be considered. Number 1 was issued
on January 23, 1957. Contributions must be not less than 8 nor exceed 72 printed pages.

INSTRUCTIONS FOR AUTHORS

Acceptance of manuscripts will be determined by the significance of new information. Priority
will be given to manuscripts by staff members. All manuscripts must be recommended by the
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Authors must adhere to the International Code of Nomenclature of Bacteria and Viruses,
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MANUSCRIPT FORM. — (1) In preparation of copy follow the 1972 CBE Style Manual,
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as footnotes will not be accepted. (3) An informative abstract must be included for all papers.
(4) A Spanish summary is required for all manuscripts dealing with Latin American subjects.
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should be sent to the Editor, Museum Publications, Natural History Museum of Los Angeles
County, 900 Exposition Boulevard, Los Angeles, California 90007.

Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

Edward Ostermeyer
Editor

A NEOGENE SECTION, NORTHEASTERN SAN CLEMENTE
ISLAND, CALIFORNIA1

By Takeo Susuki2 and Carol J. Stadum3

Abstract: A discontinuous section of Miocene, Pliocene, Pleistocene, and Re-
cent sediments overlies rhyolite 0.9 to 2.5 kilometers south of the Naval Ordinance
Test Station Pier in the area of Randall Blockhouse, northeastern San Clemente Is-
land, Los Angeles County, California. Poorly consolidated orange sandstone con-
taining the Miocene mollusk Pecten ( Amussiopecten ) lompocensis Arnold and marine
limestone lenses, grades upward into diatomaceous shale equivalent to the Miocene
Monterey Formation. Unconformably overlying the Miocene units are Pliocene bio-
clastic sediments containing the mollusks ( Pecten ) ( P .) bellus (Conrad) and Pecten
( Patinopecten ) healeyi sanclementensis n. subsp. Isolated outcrops of Pleistocene
sand, containing a molluscan fauna, correlate with the Palos Verdes Sand of the Los
Angeles Basin. An occurrence of a desmostylid tooth is reported and illustrated.

INTRODUCTION

San Clemente Island, the southernmost Channel Island off the southern California
coast, is approximately 96 km south and west of Long Beach. (Figs. 1 ,2) The elongate
island, extending northwest to southeast, is 35 km long and 2 km to 6.4 km wide.
Sediments cover the slightly tilted and gently domed block, which is composed of
lava flows, tuffs, and breccias that have been dated at about 15.7 million years (Meri-
field and others 1971). More than twenty marine terraces are exposed on the western
slope of the island, while the eastern flank rises steeply from the northwest trending
San Clemente fault.

During a visit to the island in April, 1975, an outcrop of Pliocene biosediments
was located in a discontinuous Neogene section by the junior author. The occurrence
of these strata was reported by Stadum and Susuki (1976) as containing mullusks and
brachiopods, interbedded in sediments composed predominantly of bryozoans, fora-
minifers and echinoid spines of Pliocene age and the authors’ findings were subse-
quently confirmed in a later publication by Vedder and Moore (1976). The only Chan-
nel Island known to contain similar age strata is Santa Cruz Island (Weaver and Myer
1969). An extensive field investigation was undertaken in July and November, 1975,
to determine the areal extent of the discontinuous Neogene section. The Neogene

1Review Committee for this Contribution
Warren O. Addicott
Jere H. Lipps
Edward C. Wilson

2Research Associate in Invertebrate Paleontology, Natural History Museum of Los Angeles
County, and Department of Geology, University of California, Los Angeles, California, 90024.

3Huntington Beach High School, 1905 Main Street, Huntington Beach, California, 92648.

118° 117°

Figure 1 . Index map showing general location of San Clemente Island.

1978

A Neogene Section, San Clemente Island

3

marine mollusks from northeast San Clemente Island are summarized and a new Pli-
ocene subspecies is described and illustrated.

The specimens are deposited in the paleontology collection. Department of Ge-
ology, University of California, Los Angeles.

PREVIOUS GEOLOGIC INVESTIGATIONS

The earliest publication to describe the San Clemente Island lithology was Cooper
(1865). This report stated that the “island form is that of a terraced table” and that
the rocks are composed almost entirely of basalt. He implied that the scarcity of fossils
was apparently due to lack of marine sediments. Lawson (1893) discussed the physi-
ography in detail and mentioned fossiliferous white limestone which, he thought, was
equivalent to the mainland Miocene coastal deposits. Smith (1898) recognized that
a portion of the island was composed of Miocene sandstone, shale, and limestone
with overlying sediments which were post-Pliocene in Age. Olmsted (1958) distin-
quished the lower sedimentary unit as middle Miocene and the unconsolidated sand
deposits as Pleistocene (?).

Mitchell and Lipps (1965) collected numerous Miocene marine vertebrates, pe-
lecypods, algae, and microorganisms from the Chalk Canyon area (Horse Cove) and
two mid-island basins. They stated that an interval of geologic history “from about
the end of the middle Miocene to about 500,000 years or so ago” is missing. Ocean-
ographic surveys by Ridlon (1968; 1969; 1972) found middle Miocene sedimentary
rocks, post-Miocene sediments of undetermined age, and Holocene surficial sediments
offshore near Eel Point. Merifield and others (1971) emphasized the petrography and
structure and volcanic rocks on the central part of the island, and noted the presence
of Miocene sediments and Quaternary beach sand. Most recently, Vedder and others
(1975), and Vedder and Moore (1976) discussed Miocene and Pliocene biostratig-
raphy and paleontology of the island. These publications did not include the Neogene
sediments south of Randall Blockhouse in sections RS-2 and RS-3 of this paper.

STRATIGRAPHY

Introduction

The authors have used abbreviations for four sections of Neogene sediments ex-
posed near Randall Blockhouse, an observation structure and lighthouse about 900
m south of NOTS (Naval Ordinance Test Station) Pier. Neogene sediments overlie
rhyolite and are exposed in gullies northwest, west and south of Randall Blockhouse.
The letters “RS” refer to “Randall Section”, with the number indicating the location
of outcrop (Fig. 3).

RS-1 = Pliocene sediments in roadcut above Randall Blockhouse and a ravine
just to the west of the roadcut. (includes localities 6316, 6318 and 6320).

RS-2 = Miocene, Pliocene and Pleistocene in a steep gully about 213 m south of
Randall Blockhouse, (includes localities 6321 and 6327).

4

Contributions in Science

No. 299

Figure 2. Index map of San Clemente Island showing some of the prominent landmarks.

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A Neogene Section, San Clemente Island

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RS-3 = Miocene beds about 1525 m southeast of Randall Blockhouse, (locality
6317).

RS-4 = Miocene strata about 1585 m north and slightly west of NOTS Pier.

Miocene Sandstone

The base of the Miocene sedimentary sequence is a coarse, poorly-consolidated,
unfossiliferous, dark-gray volcanic sandstone overlying rhyolite flow lava. The basal
sandstone is best developed at RS-3 (Fig. 6) with a maximum thickness of approxi-
mately 1.2 m which thins northward to about 0.3 m at RS-2, within a distance of
about 660 m. This basal sandstone was not recognized by Vedder and Moore (1976)
from the NOTS Pier area. They reported shale and claystone lying unconformably on
the rhyolite.

Grading upward from the basal volcanic sandstone is a prominent marker bed
of loosely-consolidated to well-indurated, medium to coarse grained, fossiliferous,
orange sandstone. This unit is best exposed at RS-3 (4.5 m thick), where the basal
1.2 m has abundant Pecten {Amussiopecten) lompocensis Arnold (Figs. 12-17). Echi-
noid spines, rare gastropod nodes similar to those of the genus Trophon, marine mam-
mal bones and teeth, shark teeth, and fish bones occur commonly throughout the
sandstone. At section RS-4, where the orange sand is approximately 1.5 m thick, only
a few diagnostic fragments of P. (A.) lompocensis were found. Other equivalent lo-
calities were reported by Olmsted (1958) from the central island area, including his
locality F-3 which contains fragments of Chlamys ( Lyropecten ) crassicardo (Conrad),
and his locality F-2 from which he reported “abundant shell fragments, chiefly pec-
tinids, and scattered mammalian bones.” Vedder and Moore (1976) reported Lyro-
pecten crassicardo (Conrad), Amussiopecten cf. A. vanvlecki (Arnold), and Crassos-
trea cf. C. freudenbergi (Hertlein and Jordan) in equivalent beds from the China
Canyon section, USGS loc. M6505.

Near the base of the orange sandstone are dense limestone lenses containing
Ostrea sp. with other unidentifiable shell fragments. These lenses have been observed
on the north wall of the most southerly exposure at RS-3, and near the pier at Wilson
Cove. Olmsted (1958) reported limestone at his locality F-3, overlying “gritty an-
desitic sandstone about 2.5 m thick that in turn overlies a flow of platy-jointed an-
desite.” Calcarenite deposits overlying andesite flow in China Canyon (Vedder and
Moore 1976) appear to be similar to limestone lenses at RS-3 and Wilson Cove. Bram-
lette (1946) established limestone “reef” structures at the base of the Monterey For-
mation, generally as a boundary between lower and middle Miocene sediments. Sim-
ilar middle Miocene structures are found locally in the Palos Verdes Hills areas
(Bramlette 1946) and in southwestern Orange County (Morton and others 1974).

The authors collected a large, shattered desmostylid tooth (Figs. 27, 28, 29) from
a grayish sandstone (loc. 6363) about a meter above beds containing P. (A.) lom-
pocensis and stratigraphically below the silty diatomaceous shale. The tooth was ex-
amined by L. G. Barnes, Los Angeles County Museum of Natural History, who sug-
gested that the specimen could be Paleoparadoxia tabatai, whose characteristic
cingulae have been worn off (personal communication, Mar. 24, 1977).

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Miocene Diatomaceous Shale

The diatomaceous shale member is probably the most widespread sedimentary
unit within the area investigated. Vedder and others (1975) and Vedder and Moore
(1976) have correlated this unit with the middle Miocene Monterey Formation, locally
found in the Palos Verdes Hills, San Joaquin Hills, and Capistrano syncline. The
representative section is exposed at RS-3 with a thickness of about 18 m and con-
formably overlying orange sand which becomes finer and grades upward into diato-
maceous shale. The shale is interbedded with several layers (about 7 cm thick) of
gray, finely laminated chert with abundant fish scales. Above the chert layers, four
phosphate nodule layers (each about 2.5 cm thick) are intercalated with the shale. A
bluish, unaltered vitric ash layer (15 cm thick) occurs near the top of the unit. Fish
remains, shark teeth, and marine mammal bones and teeth are common through the
shale.

Elsewhere, the shale extends north and south of NOTS Pier, and is exposed at
RS-2. More extensive sections, as much as 90 meters in thickness (Mitchell and Lipps
1965), are exposed on the southern part of the island. Vedder and Moore (1976)
extensively examined the Horse Cove and China Canyon diatomaceous deposits,
which they suggested resemble parts of the Monterey Shale exposed at Newport Bay
and in the Capistrano syncline.

The depositional sequence of orange sand with limestone lenses overlain by dia-
tomaceous shale containing chert, phosphate nodules, and vitric ash layers is equiv-
alent to the depositional sequence of middle Miocene units in the El Toro and San
Joaquin Hills area of Orange County (Morton and others 1974; Stadum 1975).

Pliocene Biogenic Sediments

The Pliocene bioclastic outcrops appear to be restricted to the eastern flank of
the central island region. The basal bed is an un sorted, thin, pebble to cobble con-
glomerate, composed of volcanic clasts uncomformably overlying the Miocene dia-
tomaceous shale. A complete section, exposed in a small ravine several hundred me-
ters south of RS- 1 , is approximately 30 m thick. Although the bottom and top contacts
are not visible, a better exposed section is in a roadcut to Randall Blockhouse (RS-
1) (Fig. 9). Pliocene sediments are also present at Lemon Tank Reservoir (Vedder
and Moore 1976) unconformably overlying Miocene sandstone on the northwest wall.
Vedder and Moore (1976) collected well-preserved P. bellus, P. healeyi, moulds of
minute gastropods Alvania and Bittium, and other fossils from a truncated calcarenite
remnant at USGS loc. M6501 at Lemon Tank Reservoir. About one hundred m east
of USGS loc. M6501, the authors found mixed Pliocene-Pleistocene sands containing
the following species: Glycymeris ( Axinola ) grewingki Dali, Cyclocardia cf. C. ven-
tricosa Gould, Pseudocardium sp., Calliostoma cf. C. gemmulatum Carpenter, Cal-
liostoma cf. C. annulatum Martyn, Architectonica cf. A. nobilis Roding, Haliotis sp.,
Opalia (O.) xvroblexvskyi (Mdrch), Terebratalia cf. T. occidentalis Dali, Laqueus van-
couveriensis diegensis Hertlein and Grant, and Paracyathus cf. P . stearnsii Verrill.

Biogenic detritus and shells compose 95% of the gently dipping, massive, coarse
Pliocene strata. Where firmly cemented, the chalky-colored biolith forms noticeable

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7

outcrops with rounded, wave-cut cavities (Fig. 1 1) greater than a meter in diameter.
There are two significant conglomerate beds (Fig. 10), with a thickness of 0.30 to
1.0 M composed of dacite, andesite, and rhyolite clasts.

The poorly-sorted bioclastic sediments are predominantly bryozoans, foramini-
fers, and echinoid spines. Within this unit, there are four fossiliferous layers con-
taining the Pliocene guide fossil Pecten ( P .) bellus (Conrad) and a new subspecies
of Pecten ( Patinopecten ) healeyi Arnold. Other species include Pecten cf. P. ( P .)
lecontei Arnold, Chlamys (C.) opuntia Dali, Pododesmus macroschisma Deshayes,
Mytilus sp., Anomia peruviana d'Orbigny, Hinnites sp., Opalia ( O .) varicostata
Steams, Epitonium ( Nitidiscala ) cf. E. ( N .) indianorum (Carpenter), and distinctive
layers of the brachiopods (Fig. 8) Laqueus californianus Koch, L. vancouveriensis
diegensis Hertlein and Grant, and Terebratalia hemphilli Dali. All invertebrates are
well-preserved. Brachiopods and pelecypods generally are articulated and faintly re-
tain their original colors.

Although similar assemblages are found in Pliocene marine deposits throughout
southern California (Pico Member of the Fernando Formation in the Ventura and Los
Angeles Basins; Santa Barbara Formation in the Ventura Basin; “San Diego” For-
mation in the western Santa Monica Mountains; Fernando Formation in the Palos
Verdes Hills and Orange County; and Niguel Formation in the San Joaquin Hills) the
lithology and fossils of the San Clemente Island Pliocene strata appear to be most
similar to the San Diego Formation in coastal San Diego County. Hertlein and Grant
(1944) described the San Diego Formation as deposits containing layers of conglom-
erates and lacking fine sediments “which indicates a water depth from low tide to
possibly fifty fathoms.” They also noted the occurrence of pockets, seams, and rare
beds of nearly pure white marl in exposures of the San Diego Formation suggesting
lithologic similarity to the San Clemente Island Pliocene.

In a more recent report, Hertlein and Grant (1960) listed Pecten ( P .) bellus ,
Pecten {Patinopecten) healeyi, Chlamys (C .) opuntia, Laqueus californianus, L. van-
couveriensis diegensis, bryozoans, and echinoids from the San Diego Formation ex-
posed at Pacific Beach. The report stated that species of the brachiopod genus Laqueus
(which is abundant in San Clemente Island Pliocene sediments) “occur at moderate
depths on clear sea bottoms free of mud, and a greater number occur in warm rather
than cold water.” By comparing the similar fauna and lithology of the San Diego
Formation with San Clemente Island Pliocene sediments, the authors suggest that the
strata were deposited in a clear, sublittoral environment that was slightly warmer than
at present.

Olmsted (1958) separated the younger deposits into Older Sand (Pleistocene) and
Younger Sand (Recent). He designated the older deposits as chiefly old dunes with
beach sand and lagoonal deposits. He stated that the distinction between the flat-lying
sandy Miocene beds overlain by older Pleistocene sand deposits “is difficult princi-
pally in the large area centered at 7 km south-southeast of Wilson Cove.” This large
area is the general location of the gently dipping Pliocene bioclastic sediments
(RS-1, RS-2), which may be the “flat-lying sandy Miocene beds” mentioned by
Olmsted.

Contributions in Science

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Pleistocene Marine Sediments

The Pleistocene consists of gray to buff color, fossiliferous sand with a basal
conglomerate, that differs in its relationship with underlying beds from locality to
locality. At RS-1 (loc. 6320) and RS-2 (loc. 6327), Pleistocene sand unconformably
overlies Pliocene sediments; about 800 m west of Wilson Cove and at NOTS Pier,
Pleistocene sand is unconformable on Miocene shale (loc. 6326); several hundred m
south of Wilson Cove (loc. 6324), it rests unconformably upon volcanic rocks; east
of Northwest Harbor near the northeast side of the new airfield, the underlying unit
is a volcanic conglomerate of unknown age (loc. 6323). The mollusks, representing
a littoral and a sublittoral environment, are found in loosely consolidated sand except
at localities 6320, 6324, and 6327, where they occur between and attached to clasts
in the basal conglomerate. Codakia ( Epilucina ) calif ornica (Conrad) was the most
commonly collected species from each locality.

The following is a list of Pleistocene fossils collected from six localities. This
assemblage can be correlated with the late Pleistocene Palos Verdes Sand fauna of
the Los Angeles Basin.

UCLA locality 6320

Codakia ( Epilucina ) californica (Conrad)

Transenella tantilla (Gould)

Acmaea { Acmaea ) mitra Eschscholtz
Acmaea ( Collisella ) scabra Gould
Hipponix tumens Carpenter

UCLA locality 6323

Codakia {Epilucina) californica (Conrad)

Gians subquadrata (Carpenter)

Acmaea {Collisella) digitalis Eschscholtz
Acmaea (C.) pelta nacelloides Dali
Acmaea (C.) scabra Gould
Acmaea (C.) scutum Eschscholtz
Fissurella volcano Reeve
Tegula funebralis (A. Adams)

Littorina planaxis Philippi
Hipponix antiquatus (Linnaeus)

Hipponix tumens Carpenter
Serpulorbis squamigerus (Carpenter)

Bittium sp.

Amphissa versiocolor Dali
Fusinus barbarensis (Trask)

Olivella biplicata (Sowerby)

Trimusculus reticulata (Sowerby)

Ischnochiton sp.

UCLA locality 6324

Codakis {Epilucina) californica (Conrad)

Fissurella volcano Reeve

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Tegula ligulata Menke
Hipponix antiquatus (Linnaeus)

Hipponix tumens Carpenter

Thais (Stramonita ) biserialis (Blainville)

Mitra idae Melvill

Hyalina californica (Tomlin)

Conus californicus Hinds
UCLA locality 6325

Codakia (Epilucina) californica (Conrad)
Acmaea ( Collisella ) limulata Carpenter
Acmaea ( C .) scabra Gould
Acmaea (C.) scutum Eschscholtz
Haliotis cracherodii Leach
Megathura crenulata (Sowerby)
Fissurella volcano Reeve
Tegula funebralis (A. Adams)

Norrisia norrisi (Sowerby)

Littorina planaxis Philippi
Hipponix antiquatus Linnaeus
Hipponix tumens Carpenter
Serpulorbis squamigerus (Carpenter)
Bursa californica Hinds
Amphissa versicolor Dali
Mitra idae Melville
Hyalina californica (Tomlin)

Conus californicus Hinds
UCLA locality 6326

Codakia ( Epilucina ) californica (Conrad)
Eissurella volcano Reeve
Astraea ( Pomaulax ) undosa (Wood)
Tegula funebralis (A. Adams)

Hipponix tumens Carpenter

Neverita recluziana (Deshayes)

Serpulorbis squamigerus (Carpenter)

Bursa californica Hinds

Thais ( Stramonita ) biserialis (Blainville)

Mitra idae Melvill

Olivella biplicata (Sowerby)

Conus californicus Hinds
UCLA locality 6327

Codakia ( Epilucina ) californica (Conrad)
Transenella tantilla (Gould)

Acmaea ( Acmaea ) mitra Eschscholtz
Acmaea (i Collisella ) limatula Carpenter
Acmaea (C.) scabra Gould

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Acmaea (C.) scutum Eschscholtz
Fissurella volcano Reeve
Tegula funebralis (A. Adams)

Hipponix antiquatus (Linnaeus)

Mitrella carinata gausapata (Gould)

Thais ( Stramonita ) biserialis (Blainville)

Olivella biplicata (Sowerby)

Conus californicus Hinds
Ischnochiton acrior Carpenter

Pleistocene Non-Marine Sediments — Recent Dune Sand

Loose, wind-blown sand accumulations cover much of the northern half of the
island and in many places contain calcareous and siliceous root sheaths or rhizocon-
cretions (Fig. 5). Johnson (1967) suggested that all the Channel Islands had supported
more vegetation during the early Holocene than at the present. The authors found
evidence of calcareous root sheaths forming around living sand-dune plants. Opaline
silica seams were common within the sand deposits west of NOTS Pier and near the
old airfield (Fig. 3). Numerous Indian midden remains and artifacts were observed
in the dune sand.

Olmsted (1958) stated that the maximum thickness of the active and recently
active sand dunes is approximately 19 meters, and that the older sand deposits are as
much as 40 meters thick west of Wilson Cove (loc. 6326).

PALEONTOLOGY

Description of New Subspecies
Class BIVALVIA Linne
Order PTERIOIDA Newell
Family PECTINIDAE Rafinesque

Type (by subsequent designation, Schmidt 1818): Ostrea maxima Linne 1758.

Subgenus Patinopecten Dali 1 898
Type (original designation): Pecten caurinus Gould 1850.

Pecten ( Patinopecten ) healeyi Arnold sanclementensis new subspecies

Genus Pecten Muller 177 6

Diagnosis. — Distinguishable by the much finer and greater number of closely
and equally spaced ribs on the left valve; ribs narrow, flat, interspaces shallow and
narrow on right valve; right valve gently but evenly convex; left valve slightly convex
with anterior and posterior margins flat to somewhat concave; poorly developed rib-
bing on umbonal area of both valves.

Description. — Shell inequivalve, wider than high, right valve slightly more con-
vex than left valve; Right valve — slightly but evenly convex; margins adjacent to ears
ornamented by numerous fine radial ribs prior to beginning of primary ribs; approx-
imately thirty, low rectangular, irregular radial ribs separated by narrower, primary

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1

shallow interspaces; primary ribs become dichotomus, separated by narrower, ex-
tremely shallow secondary interspaces and about twenty percent become trichotomus;
primary and secondary interspaces become about equal in width along the outer mar-
gin; surface of shell ornamented by numerous, closely spaced fine concentric growth
lines becoming more prominent along the margin; hinge line about one-half the shell
length; anterior ear slightly longer than the posterior ear; anterior ear separated from
body of shell by a narrow, shallow sulcus, a wide fold immediately dorsal to the
sulcus, fine radiating lines most prominent adjacent to the dorsal margin of fold,
growth lines sinuous; posterior ear with narrow sulcus, numerous fine radiating lines
with oblique growth lines. Left valve — slightly convex with anterior and posterior
margins flat to slightly concave; approximately thirty to thirty-two ribs, low, narrow
and sharp, closely but equally spaced; fine secondary riblets occur between primary
ribs; shell surface ornamented by minute concentric growth lines which become bun-
dled and prominent along the margin of shell; anterior ear ornamented with fine,
slightly sinuous growth lines crossed by radiating lines, a broad shallow sulcus extends
from the beak through the middle part of the ear; posterior ear characterized by coarse
radiating lines crossed by fine oblique growth lines.

Syntypes. — UCLA Invert. Paleo. Coll. cat. nos. 38790 (right valve), 38791 (left valve).
Locality of Syntypes . — UCLA Invert. Paleo. Coll. loc. no. 6316.

Dimensions of Syntypes. — Right valve - height - 170 mm

- length - 190 mm
Left valve - height - 137 mm

- length - 150 mm

Paratypes. — UCLA Invert. Paleo. Coll. cat. nos. 38792 (right valve), 38793 (left valve).
Locality of Paratypes. — UCLA Invert. Paleo. Coll. loc. no. 6316.

Dimension of Paratypes. — Right valve - height - 148 mm

- length - 172 mm
Left valve - height - 150 mm

- length - 155 mm

Remarks. — Numerous specimens of P. ( P .) healeyi from localities throughout
southern California were carefully examined. Considerable variation was observed
within a given locality, although none are closely similar to the San Clemente Island
forms in which the left valve shows a greater range of variation than does the right
valve. The variable features of the left valve were also discussed by Vedder and Moore
(1976) and they considered that the smooth umbonal form may possibly represent a
new species. It is quite reasonable to suspect that the specimens from San Clemente
Island may represent an extreme case of maximum variation within the species. How-
ever, the specimens from San Clemente Island differ from other described species and
display very little variation within the localities, therefore a new subspecies is estab-
lished. This subspecies is named for San Clemente Island.

PECTEN (AMUSSIOPECTEN) LOMPOCENSIS Arnold is common in the basal
part of the orange sandstone exposed at section RS-3, loc. 6317. Many of the speci-

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mens are found articulated, but complete specimens are difficult to collect. Each shell
is fractured and it is necessary to coat the surface with an adhesive, prior to removing
it from the coarse sandstone matrix.

P. (A.) lompocensis is often confused with Pecten ( Amussiopecten ) vanvlecki
Arnold, (Arnold 1907) and the two species are not readily separable unless the external
and internal surfaces are well preserved. However, there are several helpful distin-
guishing features, which include the development of the external radial ribs and the
spacing of the internal ribs. On the right valve of P. (A.) lompocensis, the external
radial ribs are poorly defined and the ribs are paired internally. On the left valve, the
external ribs are better developed and the internal ribs are not paired but equally spaced
with the interspaces. In P. (A.) vanvlecki, the external ribs on both valves are better
developed but internally, the paired ribs occur on the left valve and they are equally
spaced in the right valve.

Masuda (1971), in his study of Amussiopecten, stated that P. (A.) vanvlecki is
easily distinguishable from the frequently confused species P. (A.) lompocensis by
"its rather thick shell, squarish, low, flatly round-topped radial ribs tending to become
obsolete towards the ventral and lateral margins, paired internal ribs developed at
lower part and larger and angulate auricles.” Masuda (1971) did not make any dis-
tinction as to which valve is characterized by the paired internal ribs.

The occurrence of fragmentary specimens of Amussiopecten cf. A. vanvlecki is
reported by Vedder and Moore (1976) from China Canyon in the southern end of San
Clemente Island in association with Lyropecten crassicardo (Conrad) and Crassostrea
cf. C . freudenbergi (Hertlein and Jordan), USGS loc. M6505. According to Vedder
and Moore (1976), these fossils are imbedded in a fine-to coarse-grained calcarenite
which differs considerably in lithology from the orange sandstone in which P. (A.)
lompocensis occurs so commonly (UCLA loc. 6317, which apparently has not been
visited by previous investigators).

CONCLUSIONS

Unusually well-preserved marine mollusca are found in Neogene sedimentary
units on San Clemente Island. Miocene sands contain pecten beds and oyster lenses
which suggest a shallow, near-shore environment. Diatomaceous shales, overlying
these sands, have been correlated with the coastal Monterey Formation and are evi-
dence of deep submergence during the middle Miocene. These Miocene deposits
appear to have accumulated in the same depositional basin as the Monterey Formation
in the southeastern Los Angeles Basin and represent a comparable environment.

The overlying Pliocene strata contain abundant brachiopods, bryozoans, and pec-
tens with conglomerate layers which are indicative of shallow water and uplift of the
island block. The small fauna includes diagnostic Pliocene species which correlate
with the Pecten (P.) healeyi zone (upper Pliocene) throughout southern California.

Abundant collections of Pleistocene littoral and sublittoral mollusks in sand de-
posits and among volcanic boulders continue to denote elevation of the island. The
formation of marine terraces and the deposition of dune sand with rhizoconcretions
mark the emergence of the island during the Holocene.

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Figure 3. Map of northern and part of central San Clemente Island showing fossil localities
and RS-sections.

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1 1 -5

Figure 4. A comparison of generalized stratigraphic columns, Randall Blockhouse Area.

Figures 5-8. (5.) Rhizoconcretions in post-Pleistocene sands. (6.) Miocene section at RS-3, the
darker unit just above the collectors is the orange sandstone. (7.) Pecten ( Amussiopecten )
lompocensis occurring in the orange sandstone, UCLA loc. 6317. Icepick points to a bone.
(8.) Brachiopod bed exposed in roadcut at RS-1.

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II

Figures 9-1 1 . (9.) Soil-covered Pliocene sediments exposed in roadcut just above Randall Block-
house, section RS-1. (10.) Characteristic conglomerate layer (middle of photograph) in Pliocene
strata. (11.) Rounded, wave-cut cavities in Pliocene strata at RS-2.

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Figures 12-17. (12.) Pecten ( Amussiopecten ) lompocensis Arnold, left valve, external view,
height - 64 mm. length - 64 mm. UCLA cat no. 38788. UCLA loc. 6317. (approx. X 7/10).
(13.) Pecten ( Amussiopecten ) lompocensis Arnold, left valve, internal view, height - 70 mm.
length - 70 mm. UCLA cat no. 38896. UCLA loc. 6317. (approx. X 3/5). (14 .) Pecten (Amus-
siopecten) lompocensis Arnold, right valve, internal view, height - 44 mm. length - 41 mm. UCLA
cat no. 38895. UCLA loc. 6317. (approx, x 4/5). (15). Pecten (Amussiopecten) lompocensis
Arnold, right valve, external view, height - 95 mm. length - 100 mm. UCLA cat no. 38789, UCLA
loc. 6317. (approx. X 1/2). (16 .) Pecten (Amussiopecten) lompocensis Arnold, right valve, ex-
ternal view, height - 64 mm. length - 64 mm. UCLA cat no. 38788. UCLA loc. 6317. (approx.
X 7/10). (17 .) Pecten (Amussiopecten) lompocensis Arnold, right valve, internal view, height -
44 mm. length - 47 mm. UCLA cat no. 38897. UCLA loc. 6317. (approx, x 4/5).

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Contributions in Science

Figures 18-19. (18.) Pecten {Patinopecten) healeyi sanclementensis n. subsp. Syntype. right
valve, height - 170 mm. length - 190 mm. UCLA cat no. 38790. UCLAloc. 6316. (approx. X 2/5).
(19 .) Pecten {Patinopecten) healeyi sanclementensis n. subsp. Syntype. left valve, height - 137
mm. length - 150 mm. UCLA cat no. 38791. UCLA loc. 6316. (approx, x Vz).

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19

21

Figures 20-21. (20 .) Pecten (Patinopecten) healeyi Arnold. Hypotype. right valve. Figured by
Hertlein & Grant. 1972 (text-fig. 9A, p. 184). height - 122 mm. length - 130 mm. UCLA cat no.
1950. Pacific Beach, San Diego, California (approx. X V2) compare with fig. 18. (21.) Pecten
( Patinopecten ) healeyi Arnold. Hypotype. left valve. Figured by Hertlein & Grant. 1972 (text-
fig. 9B, p. 184). height - 122 mm. length - 130 mm. UCLA cat no. 1950. Pacific Beach, San
Diego, California, (approx. X V2) compare with fig. 19.

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Figures 22-26. (22.) Opalia ( Opalia ) wroblewskyi (Morch). ab-aperture view, height - 81 mm.
width -22 mm. UCLA cat no. 38798. UCLAloc. 6322. (approx. X 9/10). (23.) Same specimen as
fig. 22. aperture view, (approx, x 9/10). (24 .)Pecten (Pecten) cf. P. (P.) lecontei Arnold, right
valve, height - 60 mm. length - 67 mm. UCLA cat no. 38787. UCLA loc. 6316. (approx, x 9/10).
(25 .) Pecten ( Pecten ) bellus (Conrad), right valve, height - 39 mm. length - 43 mm. UCLA cat
no. 38786. UCLA loc. 6316. (approx, x 9/10). (26 .) Chlamys (Chlamys) opuntia (Dali), right
valve, height -66 mm. length -66 mm. UCLAcatno. 38797. UCLAloc. 6316. (approx, x 9/10).

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Figures 27-29. (27.) Large desmostylid tooth, occlusal view. UCLA cat no. 57531. UCLA loc.
6363. (approx. X 2) length - 33 mm. width - 22 mm. (28.) Same specimen as fig. 27. end view,
(approx. X 2) height - 44 mm. (29.) Same specimen as fig. 27. labial view, (approx. X 2).

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ACKNOWLEDGMENTS

The authors are indebted to Herb Beltran, Foreman Rigger, Naval Undersea
Center (NUC), San Clemente Island, who arranged for the initial visit to the island
and provided invaluable aid in camp and in the field. Jack Harris, Range Engineer
NUC, arranged for subsequent visits to the island. Those, who were responsible for
permitting research to be conducted on the island are Lt. Commander W. G. Kay,
Officer-in-charge, San Clemente Island, NUC; Commander Timothy C. Kelley, CEC
USN, Public Works Officer, NUC; and Jan K. Larsen, Wildlife Biologist, Natural
Resources Program, NUC.

The following people contributed significantly in the field: Carl Westendarp,
NUC; Marvin Rohrs and Lorna Ross, Natural History Foundation of Orange County;
Mark Beck, Bill and John Beltran, Danny Cook, Richard Ebel, Alan Leach, John
Lemo, Lorry Partridge, Cindy Phelps, and Roger Zachery, Huntington Beach High
School. Lawrence G. Bames, Earth Science Division, Los Angeles County Museum
of Natural History examined the desmostylid tooth. The manuscript was kindly read
and criticized by Willis P. Popenoe and Paul M. Merifield, Department of Geology,
University of California, Los Angeles. Recognition is given to Edward C. Wilson,
Earth Science Division, Los Angeles County Museum of Natural History, Warren O.
Addicott, Department of Paleontology and Stratigraphy, United States Geological
Survey, Menlo Park, California, and Jere H. Lipps, Department of Geology, Uni-
versity of California, Davis, California for their critical review.

All illustrations and photographs were prepared by the authors.

FOSSIL LOCALITY REGISTER

The following locality numbers are those of the paleontology collection, De-
partment of Geology, University of California, Los Angeles. They all were plotted
on the U.S. Geological Survey topographic quadrangle of San Clemente Island North
(7.5 min., 1950), unless otherwise indicated.

6316 - Sediments containing Pecten ( P .) bellus and Pecten (P.) healeyi sanclementensis n.
subsp. exposed in both sides of a roadcut, just before last turn to the end of road to Randall
Blockhouse (RS-1), approx. 792 m S 42° E of NOTS Pier. (Upper Pliocene).

6317 - Loosely consolidated orange sandstone bed containing Pecten (A.) lompocensis exposed
in a narrow, steep canyon approx. 1524 m S 35° E of NOTS Pier and about 1097 m N 58°
E of BM-820 (RS-3). (Middle Miocene)

6318 - Well-indurated chalky-colored sediments cropping out on the south side of a narrow
ravine containing Pecten ( P .) bellus, about 213 m west of Randall Blockhouse (RS-1),
(Upper Pliocene)

6320 - Pleistocene fossils attached to boulders and in the sand between the conglomerate on the
southside of a narrow ridge about 427 m S 13° W of Randall Blockhouse (RS-1) and approx.
700 m S 18° W of NOTS Pier. (Upper Pleistocene)

6321 - Steep, small gully south of Randall Blockhouse (RS-2), Pliocene section exposed un-
conformably overlying Miocene shale and underlying Pleistocene conglomerate, about 213
m south of Randall Blockhouse. (Upper Pliocene)

6322 - Fossils collected from disturbed (by grading) beds overlying the brownish-orange sand-
stone on the northwest wall of Canyon (Lemon Tank) just below the top, 365 m east of

1978

A Neogene Section, San Clemente Island

23

Hill 1185, U.S. Geological Survey topographic guadrangle of San Clemente Island Central
(7.5 min., 1950). (Upper Pliocene with some Upper Pleistocene)

6323 - Fossils collected from north of northeast end of new airfield, halfway down the slope
between upper road to the first wide, flat terrace, 1280 m S 3° E of stream entering North-
west Harbor. (Upper Pleistocene)

6324 - Pleistocene along the top edge of sea wall in conglomerate, overlying volcanic rocks,
about 455 m S 65° E from the end of Wilson Cove Pier. (Upper Pleistocene)

6325 - Pleistocene fossils in conglomerate and loose sand overlying Miocene shale, directly
behind building by NOTS Pier. (Upper Pleistocene)

6326 - Fossiliferous Pleistocene bed in a shallow cut, lying unconformably on Miocene shale,
365 m S 68° E of Harbor triangulation station 648. (Upper Pleistocene)

6327 - Pleistocene fossils in conglomerate unconformably overlying Pliocene sediments in a
gully 183 m due south of Randall Blockhouse (RS-2). (Upper Pleistocene)

6363 - A large desmostylid tooth collected from a gray sandstone approximately one hundred
m south of UCLA loc. 6317, stratigraphically about a meter above beds containing Pecten
( Amussiopecten ) lompocensis .

LITERATURE CITED

Arnold, Ralph. 1906. Tertiary and Quaternary pectens of California. U.S. Geol. Surv. Prof.
Paper 47: 92-93; pi. 28, figs. 1, 2, 3.

1907, Fossil mollusks from California. Smithsonian Misc. Coll. 50: 428-429; pi.

Lin, fig. i, 2.

Bramlette, M. N. 1946. Monterey Formation of California and the origin of its siliceous rocks.
U.S. Geol. Surv. Prof. Paper 212: 55 p.

Cooper, J. G. 1865. Geology of California, Sect. V. Islands off the coast of southern California.
Geol. Surv. of California. 1: 182-186.

Hertlein, L. G. and U. S. Grant. 1944. The geology and paleontology of the marine Pliocene
of San Diego, California. Part I. Geology, San Diego Soc. Natur. Hist. Mem. II: 72 p.

1960. The geology and paleontology of the marine Pliocene of San Diego, California.

Part 2a. Paleontology. San Diego Soc. Natur. Hist. Mem. II: 73-133; pis. 19-26.

1972. The geology and paleontology of the marine Pliocene of San Diego, California.

(Paleontology: Pelecypoda). San Diego Soc. Natur. Hist. Mem. II. Part 2b: 143-409; pis.
27-57.

Johnson, D. L. 1967. Caliche on the Channel Islands. California Div. Mines & Geol. 20 (2):
151-158.

Lawson, A. C. 1893. The post-Pliocene diastrophism of the coast of southern California. Bull.
Dep. Geol. Univ. California 1 (4): 135-139.

Masuda, Koichiro. 1971. Amussiopecten from North America and northern South America.

Palaeontological Soc. Japan Trans, and Proc. 84: 205-224: pis. 25, 26.

Merifield, P. M., D. L. Lamar and M. L. Stout. 1971. Geology of central San Clemente
Island, California. Geol. Soc. Amer. Bull. 82 (7): 1989-1994.

Mitchell, Edward D., Jr. and Jere H. Lipps. 1965. Fossil collection San Clemente Island.
Pacific Discovery. 18 (3): 2-8.

Morton, Paul K., William J. Edgington and Donald L. Fife. 1974. Geology and en-
gineering geologic aspects of the San Juan Capistrano Quadrangle, Orange County, Cali-
fornia. California Div. Mines & Geol. Spec. Rep. 112: 12, 48-49.

Olmsted, F. H. 1958. Geologic reconnaissance of San Clemente Island, U.S. Geol. Surv. Bull.
1071-B: 55-68.

24

Contributions in Science

No. 299

Ridlon, J. B. 1968. San Clemente Island Rocksite Project. Offshore Geology. Part 1. Detailed
survey off Eel and Lost Points area. Dep. of the Navy, Naval Weapons Center, China Lake,
California: 79 p.

1969. San Clemente Island Rocksite Project. Offshore Geology. Part II. Reconnais-
sance survey around the island. Dep. of the Navy Weapons Center, China Lake, California:
132 p.

1972. Pleistocene-Holocene deformation of the San Clemente Island crustal block,

California. Geol. Soc. Amer. Bull. 83 (6): 1831-1844.

Smith, W. S. T. 1898. A geological sketch of the San Clemente Island, U.S. Geol. Surv. 18th
Ann. Rep. Part 2: 459-496.

Stadum, Carol J. 1975. “Paleontological Sites” Construction of regional wastewater facilities,
Environmental Impact Report for Aliso Water Management Agency. Culp/Wesner/Culp,
Clean Water Consultants: III- 13- 19.

Stadum, Carol J. and Takeo Susuki. 1976. The discovery of marine Pliocene strata on San
Clemente Island, California. Geol. Soc. Amer. Abstracts with programs. 8 (3): 411; Feb.
1976.

Vedder, J. G., L. A. Beyer, A. Junger, G. W. Moore, A. E. Roberts, J. C. Taylor and
H. C. Wagner. 1975. Preliminary report on the geology of the continental borderlands of
southern California, U.S. Geol. Surv. Map MF-624: 9 maps.

Vedder, J. G. and Ellen J. Moore. 1976. Paleoenvironmental implications of fossiliferous
Miocene and Pliocene strata on San Clemente Island, California: p. 107-135. In D. G.
Howell ed. Aspects of the geologic history of the California continental borderland. Pacific
Section A.A.P.G. Misc. Pub. 24.

Weaver, D. W. and G. L. Myer. 1969. Stratigraphy of northeastern Santa Cruz Island,
p. 158-172. In D. W. Weaver and others. Geology of the northern Channel Islands, south-
ern California borderland. (Los Angeles, California) A.A.P.G. & S.E.P.M. Pacific Section.

Accepted for publication March 30, 1977.

f,

NUMBER 300
JUNE 16, 1978

A SYSTEMATIC REVIEW OF THE MEXICAN FROG
RANA SIERRAMADRENSIS TAYLOR

By Robert G. Webb

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

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Printed in the United States of America by Chapman’s Phototypesetting on 70# Patina Book

A SYSTEMATIC REVIEW OF THE MEXICAN FROG
RAN A SI ERR A MA DRENSIS TAYLOR1

By Robert G. Webb2

Abstract: The variation of Rana sierramadrensis Taylor is discussed. A pre-
viously unreported secondary sexual characteristic is the presence of white spinous
excrescences on the non-enlarged thumbs, chest and lower jaw of males. The tadpoles
are described. Rana sierramadrensis is compared with R. sinaloae Zweifel. The former
is retained in the R. palmipes species group, while the latter is transferred to the R.
tarahumarae group, which is proposed for the inclusion of the Mexican species, R.
tarahumarae , R. pustulosa, R. sinaloae , R. pueblae, and R.johni. Rana sierramadren-
sis is confined to the Sierra Madre del Sur in the Mexican states of Guerrero and Oaxaca,
and seems to be most closely related to Rana maculata.

INTRODUCTION

Rana sierramadrensis is an upland tropical species in southern Mexico that is
little known except for a few published locality records. My interest in R. sierra-
madrensis is a by-product of a study of frogs of the R. pustulosa-tarahumarae com-
plex and R. sinaloae in western Mexico. Zweifel included both R. sierramadrensis
and R. sinaloae in the R. palmipes species group (1954), and the two species R.
pustulosa and/?, tarahumarae in the/?, boylei group (1955). In view of speculation
that /?. sinaloae is related to the /?. pustulosa-tarahumarae complex, most available
specimens of/?, sierramadrensis were examined in order to determine its relationship
to /?. sinaloae .

The “ Rana tarahumarae group” is proposed for the first time to accommodate
the Mexican species/?, tarahumarae, R. pustulosa, R. sinaloae, R. pueblae and/?.
johni. Biochemical data (Case 1976 and personal communication) suggest that /?.
tarahumarae (only Mexican species studied by Case) is not closely related to the two
United States members of Zweifel’s /?. boylei group (1955), /?. boylei and/?, muscosa.
Those two species also differ from/?, tarahumarae and the other Mexican species in
having two metatarsal tubercles (instead of one), and the larvae having four or more
lower rows of teeth (instead of three), and in lacking marginal teeth.

Specimens examined are deposited in the following collections: American Mu-
seum of Natural History (AMNH); Field Museum of Natural History (FMNH); Natural
History Museum of Los Angeles County (LACM); University of Kansas Museum of

’Review Committee for this Contribution
Robert L. Bezy
Roy W. McDiarmid
John W. Wright
Richard G. Zweifel

2Research Associate in Herpetology, Natural History Museum of Los Angeles County, and De-
partment of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968

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Contributions in Science

No. 300

Natural History (KU); The Museum, Michigan State University (MSU); Texas Co-
operative Wildlife Collection, Texas A&M University (TCWC); University of Illinois
Museum of Natural History (UIMNH); University of Michigan Museum of Zoology
(UMMZ); and, National Museum of Natural History (USNM). I thank Richard G.
Zweifel for a kodachrome slide and a black-and-white photograph of Rana maculata
here reproduced in figure 5.

Rana sierramadrensis Taylor

Rana sierramadrensis has always been considered a distinct, monotypic species.
There are no known synonyms of R. sierramadrensis . In the original description,
Taylor (1939:397-398) listed six specimens (3963A, 3963B, 6565, 6566, 6567, and
6568), designating 3963B as the holotype and the remainder as paratypes; these num-
bers refer to the Edward H. Taylor-Hobart M. Smith private collection (EHT-HMS).
Later, Taylor (1944:140) reiterated data about the types. Of the six specimens in the
type series, four were subsequently sent to the FMNH and two were donated to the
UIMNH. The type material is briefly discussed below.

The holotype (Fig. 1, formerly EHT-HMS 3963B) from “near Agua del Obispo,
between Rincon and Cajones, Guerrero, July 1, 1932“ is now FMNH 100038. It is
a female 88.2 mm SVL and was described in some detail by Taylor (1939:398-399).
The other large frog (formerly EHT-HMS 3963 A) with the same data of collection
as the holotype is UIMNH 27053 (Smith, Langebartel, and Williams 1964:32); a pho-
tograph of this paratype, 69.7 mm SVL, was published in the original description
(Taylor 1939: PI. XXIX, Fig. 1). Four small paratypes do not exceed 40 mm SVL.

Figure 1. Holotype of Rana sierramadrensis, FMNH 100038, from near Agua del Obispo,
between Rincon and Cajones, Guerrero, Mexico.

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A Systematic Review of the Mexican Frog

3

One of them (formerly EHT-HMS 6565 and UIMNH 27054), 38.9 mm SVL from
“about 9 km. southwest of Mazatlan, Guerrero (km. 337), July 21, 1936,” is now
USNM 139724 (Cochran, 1961:76). A small paratype (formerly EHT-HMS 6566),
33.5 mm SVL, from “near Agua del Obispo (km. 350-351), July 24, 1936,” is now
cataloged as FMNH 102202. The other two paratypes (formerly EHT-HMS 6567 and
6568), 37.0 and 28.8 mm SVL, both from “Agua del Obispo, August 1, 1936,” are
now cataloged, respectively, as FMNH 102201 and FMNH 103917. Museum locality
data associated with some paratypes are slightly different from the localities quoted
(above) by Taylor (1939).

Description

Color and pattern. — The dorsal ground color is brown or bronze-brown. The
narrow, dorsolateral folds are usually slightly paler (usually buff, not white) than the
ground color, have distinct, but narrow, black, lateral borders, and extend to the sacral
region. A prominent white supralabial stripe extends from near the tip of the snout
to above the insertion of the forelimb; this white stripe is widest posteriorly (above
forelimb insertion) where it is often interrupted forming a posteriormost, white spot.
There is a black canthal stripe, black diagonal posttympanic bar, black barlike mark
on the anterior surface of the humerus at the insertion of the forelimb, and often a
black blotch or barlike mark in the groin. A posttympanic fold is absent; however,
the region is often somewhat swollen and a partial fold is discemable in AMNH
52624. Ventrolaterally the body has black markings that form an irregular-bordered,
continuous or interrupted band extending from axilla to groin. The femora have nar-
row, dorsal, dark crossbars (pale interspaces usually three to five times wider), and
a coarse pale-dark marbling (mostly dark) on the posterior surfaces. The ventral sur-
face of the legs also has coarse dark marbling resulting mostly from lateral encroach-
ment of pattern. The top of the head, back, and usually the sides of the body are
devoid of pattern; occasionally, a few small black dots occur on the back. The ventral
surface of the head-body may lack dark marks or smudging (LACM 35054; UIMNH
32444, 52783), but usually the chin-throat, chest and anterior part of the abdomen
are dark-smudged and often uniformly dark. The features of color and pattern are
most contrasting in young and subadults (Fig. 5) and tend to become obscured in the
largest frogs (Fig. 1).

Sexual dimorphism . — There seems to be no marked discrepancy in the maximum
size attained by males (100 mm SVL, KU 87284) and females (98 mm SVL, TCWC
8546). My examination of large males does not reveal vocal sacs or slits. A previously
unreported, secondary sexual character in males is the occurence of whitish, spinous,
nuptial excrescences on the thumb that, at maximal development, extend onto the
chest and edge of the lower jaw. All frogs having these spinous excrescences were
determined by dissection to be males. There is no pronounced enlargement of the
thumb in large males. A cluster of nuptial spines first appears on the joint between
the penultimate and antepenultimate phalanges. The white, spinous patch then spreads
over the inner, medial surface of the thumb. With further development small excres-
cences appear on the medial surfaces of the second and third fingers. At maximum

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No. 300

development spinous excrescences also occur on the dorsal surfaces of the first two
fingers (and part of the third finger), and on the edge of lower jaw and part of the
chest; the excrescences are more dense on the thumb joint than elsewhere (Fig. 2).
Spinous patches are not evident until males attain a body length of about 70 mm;
thereafter, however, the development and extent of the excrescences does not seem
to be directly correlated with size. The smallest male with spinous patches (covering
only proximalmost joint of thumb) is 74.2 mm SVL (KU 87280). Two other males,
both dissected, of 74.0 (UMMZ 125901) and 84.4 (KU 87283) mm SVL lack ex-
crescences. In contrast, a male of 85.8 mm SVL (UIMNH 52782) shows the most
extensive development with white, spinous excrescences on digits, chest and lower
jaw (Fig. 2). This non-correlation of size with development of excrescences would
seem to reflect a seasonal change with maximum development during the breeding
season. If so, the breeding season is prolonged since the two males showing the most
extensive development of white excrescences, UIMNH 52782 and KU 87284 of 100.3
mm SVL, were collected on 25 December 1962 and 3 June 1964, respectively.

Body proportions . — Four morphometric characters were utilized. The following
measurements (mm) were recorded with a dial calipers (abbreviations used in paren-
theses): snout to vent length (SVL); tibia (tibio-fibula) length from heel to fold of
skin on knee (TL); head length from posterior margin of tympanum to tip of snout
(HL); head width at widest point, generally at level of posterior margin of tympana
(HW). Six proportions of body parts (HW/SVL, HL/SVL, HL/HW, TL/SVL, HL/
TL, HW/TL) were plotted against SVL; the resultant data thus reflect relative differ-
ences in width of head and length of leg. Ontogenetic variation is variable depending
on the ratios utilized. Sexual dimorphism is lacking and the sexes are combined in
the analysis of variation. The data are presented in Table 1. The size-group demar-
cation (60 mm) was initially determined by the inspection of scattergrams that depicted
the most pronounced ontogenetic variation in the ratios TL/SVL, HL/TL, and HW/
TL.

In the ratio HW/SVL, ontogenetic variation seems to be negligible or the HW
increases at a slightly slower rate than the SVL. Ontogenetic variation is somewhat
more pronounced in HL/SVL with the HL increasing at a slightly slower rate than the
SVL. Heads are usually longer than broad (HL/HW exceeding 1.00). Although the
data suggest much variation in small frogs, the heads tend to become relatively more
broad with increasing size with some of the largest frogs having heads broader than
long (two males). The ontogenetic variation in HL/HW thus seems to be reflected in
the relatively slow rate of increase of HL (HW increasing at about same rate as SVL).
There is more pronounced ontogenetic variation in TL/SVL with the TL increasing
at a faster rate than the SVL; large frogs have on the average relatively longer legs
than juveniles, with their TL averaging slightly more than half the body length. On-
togenetic variation is also expressed when the ratios HW/TL and HL/TL are plotted
against SVL, with the variation most pronounced in HL/TL (owing to slow rate of
increase of HL and fast rate of increase of TL relative to that of SVL).

In comparing ratios derived from body measurements in different kinds of frogs,
it is necessary to determine the presence (if any) and degree of ontogenetic variation
in each. The comparison of ratios may be taxonomically useful only when utilizing

1978

A Systematic Review of the Mexican Frog

5

Figure 2. Adult male of Rana sierramadrensis (UIMNH 52782, 85.8 mm SVL) showing white,
spinous, excrescences on chest and lower jaw (above), and on dorsal surfaces of inner three
digits of right forelimb (below).

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Contributions in Science

No. 300

Table 1

Ontogenetic variation in six body proportions of Rana sierramadrensis. The upper
set of values is for frogs less than 60 mm S VL, the lower set for frogs 60 mm S VL or
larger. The data for each proportion are mean ± its standard error, one standard
deviation, and range (in parentheses).

Ratio

N

Variation

HW/SVL

14

0.357 ± 0.005, 0.018 (0.33-0.39)

36

0.346 ± 0.002, 0.013 (0.31-0.37)

HL/SVL

14

0.383 ± 0.004, 0.016 (0.36-0.41)

36

0.363 ± 0.003, 0.016 (0.33-0.41)

HL/HW

14

1.072 ± 0.001,0.054(0.97-1.15)

36

1.043 ±0.306,0.035 (0.94-1.12)

TL/SVL

14

0.506 ± 0.008, 0.031 (0.44-0.54)

36

0.568 ± 0.004, 0.025 (0.53-0.63)

HW/TL

14

0.708 ± 0.002, 0.057 (0.63-0.81)

36

0.610 ± 0.004, 0.026 (0.56-0.66)

HL/TL

13

0.746 ± 0.001 , 0.053 (0.67-0.84)

36

0.638 ± 0.005, 0.030 (0.58-0.68)

restricted size groups. An on-going study of the/?, pustulosa-larahiimarae complex,
for example, indicates differences in ontogenetic variation between taxa in some ra-
tios. The foregoing data for Rana sierramadrensis may be useful to future investi-
gators in subsequent comparisons with related species.

Larvae. — The larvae or tadpoles of Rana sierramadrensis, not previously de-
scribed, are discussed below and compared with larvae of sympatric and related ranid
species in Mexico. Three tadpoles (KU 87660, N = 4) from 3.3 km north San Vi-
cente, Guerrero, 920 m, collected on 8 June 1964 are assigned to R. sierramadrensis
(see below). Another tadpole included in this lot of four tadpoles is smaller (24 mm
total length) than the other three; the upper tooth rows are lacking (three lower rows
present), the lateral margin of the oral disc is emarginate, and the dark-blotched tail
fin has numerous filamentous melanophores. This small tadpole does not seem to be
representative of Rana sierramadrensis . The description is based on the three larger
larvae that exhibit the ranid features of sinistral spiracle, dextral anus, eyes more
dorsal than lateral, and a papillate fringe along the lower lip; however, the lateral
margins of the oral disc are not emarginate.

The three larvae measure 38, 41, and 50 mm in total length and correspond to
limb bud stages I and V, and paddle stage X, respectively, according to the ranid
staging system of Taylor and Kollros (in Rugh, 1962:70-71); corresponding stages of
Gosner (1960) are 25, 30, and 35. The bodies of all three larvae have obscurely
margined, broad pale dorsolateral areas. All three larvae either lack or have only
sparse pigmentation on the tail and fins. The ventral tail fin is completely devoid of
melanophores. The mostly clear dorsal fin has only a few small scattered dark pigment

1978

A Systematic Review of the Mexican Frog

7

patches (dendritic melanophores) and in some places the otherwise clear parts of the
fin contain some scattered punctate (not filamentous) melanophores. The tail mus-
culature is lightly and mostly uniformly pigmented, most dense on the dorsal half,
and lacks definitive spots-blotching. The relatively streamlined tail with the reduced
and not highly arched dorsal fin is illustrated (smallest larva of 38 mm) in figure 3D;
however, the dorsal fin seems to be slightly more arched in the two larger larvae. The
lateral margins of the oral disc are not infolded between the upper and lower tooth
rows. There are submarginal papillae along the sides of the upper and lower lips. The
fleshy flaplike lower lip with a papillate fringe seems unusually broad (especially larva
of 38 mm) and may be folded down (Fig. 3C) or up against the lower tooth rows.
The oral disc is large; the relative sizes of the oral discs of R. sierramadrensis and
the sympatric /?. pustulosa (stages X (35) and II (26), respectively; stages of Gosner,
1960, in parentheses) are compared in similar-sized larvae in figure 3A and 3B. The
tooth row formula, modified from Altig’s terminology (1970), is 7(2-7)/ 1 -4/3( 1 ) ; the
numbers between diagonal lines indicate rows of marginal teeth. The edges of the
homy beaks are pigmented and serrated. One of the larvae (41 mm) has irregular
alignment of some upper tooth row segments that number eight when counted on the
left side. Careful manipulation is required to discern the short seventh upper row of
teeth on the right side of the largest larva. The first upper tooth row is continuous,
all other upper tooth rows are separated, most by the upper beak. Larvae may have
six upper tooth rows (and perhaps eight) owing to ontogenetic and/or individual vari-

Figure 3. A, large oral disc or larva (50 mm total length) of Rana sierramadrensis (KU 87660, N =
3); B, small oral disc of larva (52 mm total length) of Rana pustulosa, compare with A; C,
mouthparts of larva (38 mm) of R. sierramadrensis; D, shape and melanophore pattern of tail of
larva (38 mm) of R. sierramadrensis.

B

D

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No. 300

ation. The A-2 gap is narrow (about equal to the width of five or six individual teeth)
or the two lateral segments are nearly juxtaposed medially. The marginal teeth (corners
of oral disc) may be scattered or arranged in rows; there are four small rows on the
right side in the largest larva. The first lower tooth row is narrowly interrupted me-
dially, the gap about equal in length to the medial gap in row A-2; the other two lower
tooth rows are continuous. The oral disc and mouthparts are illustrated in figure 3C.

The three tadpoles assigned to R. sierramadrensis are not positively identified
through direct association with metamorphosed individuals, although some such frogs
(KU 87276-80) were collected at the same locality as the larvae. The three larvae are
presumed to be of this species based on marked differences when compared with
numerous larvae of thei?. pipiens andiC tarahumarae groups from western Mexico.
See subsequent section for comparison of larvae.

Distribution

Rana sierramadrensis occurs along tropical- subtropical, foothill-montane, swilt-
flowing, often cascading, streams of the Sierra Madre del Sur in the Mexican states
of Guerrero and Oaxaca (Fig. 4). A total of 70 specimens was examined (some of
these are late-transforming frogs or are damaged and were not included in the data

Figure 4. Map of southern Mexico showing localities (solid circles) for Rana sierramadrensis
in the states of Guerrero and Oaxaca. Some localities a short distance apart share the same
symbol.

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A Systematic Review of the Mexican Frog

9

analysis). The type-locality (Agua del Obispo) and other place-names in the Chilpan-
cingo area of Guerrero were mapped by Davis and Dixon (1959).

Guerrero : near Agua del Obispo, between Rincon and Cajones (FMNH 100038,
102201-02, 103917; KU 87282; TCWC 10992; UIMNH 27053, 32442, 32444;
UMMZ 115419, 125902-N = 5, 125903; USNM 114009-12; 9 km. SW Mazatlan,
km. 337 (USNM 139724); 5.8 mi. S Mazatlan (UMMZ 115420); 3.7 mi. S Mazatlan
(UMMZ 115421); 1 mi. SE San Andres de la Cruz (UMMZ 125901, N = 4); 3.3
km. N San Vicente (KU 87276-80); 37 km. S Chilpancingo (KU 87281); Acahuizotla
(TCWC 8533-37, 8540-41, 10220-28, 26380; LACM 35054-55); 6 mi. NW San Je-
ronimito (UMMZ 125902, N = 5).

Oaxaca : 6 mi. N San Gabriel Mixtepec (MSU 10464); 6 km. NNW San Gabriel
Mixtepec (KU 87283); 12 km. NNW San Gabriel Mixtepec (KU 87284); 14.8 km.
N San Gabriel Mixtepec (KU 137539-40); Rio Jalatengo, 0.8 km. S Jalatengo (KU
137538); 5.1 km. S Jalatengo (KU 137537); 13.1 km. N Juchatengo (KU 137541);
11 km. S Chicahuaxtla (MSU 12660); 3 mi. S Putla (UIMNH 52783-84); Cacahau-
tepec, at river (UIMNH 52782); Santa Lucia, near Tehuantepec (AMNH 52624).

Comparison with Rana sinaloae

Rana sinaloae shares some features of color and pattern with R. sierramadrensis
— evidence of dark face mask bordered below by distinct white supralabial stripes;
pale dorsolateral stripes; top of head, back and sides of body mostly devoid of pattern;
narrow, dark, dorsal crossbars on femora; and black, irregular-bordered, and often
interrupted, ventrolateral bands on body (see comparison in Fig. 5). Rana sinaloae
is also similar to R. sierramadrensis in the body proportions affected by ontogenetic
variation, especially HL/SVL and TL/SVL. The two species both have heads that in
most specimens are longer than broad, and have relatively long legs (TL increasing
at faster rate than SVL).

Rana sinaloae differs from/?, sierramadrensis in having folds above and behind
the tympana, in lacking distinct black posttympanic bars, in lacking distinct black
lateral borders along the dorsolateral folds, and in having the dorsolateral folds (when
present) terminating before the groin. Perhaps most important, the breeding males of
R. sinaloae have swollen glandular thumb pads, instead of the non-enlarged thumbs
and white, spinous excrescences of males of R. sierramadrensis. Rana sinaloae shares
all these features with the other included species of the/?, tarahumarae group. There
are also corresponding differences in the larvae (see below). Rana sinaloae is con-
sidered to be a member of the Rana tarahumarae group and not a member of the
Rana palmipes species group

Comparison of Larvae

Since/?, sierramadrensis is geographically sympatric with frogs of the/?, tar-
ahumarae group (currently referred to /?. pustulosa ) and /?. pipiens group (hereafter
referred to as /?. pipiens ), larvae of any one, or two, or all three, species could be
expected to occur in a random sample. The larvae of/?, pipiens are easily distinguished
from those of/?, sierramadrensis in having a maximum of three upper tooth rows

10

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No. 300

Figure 5 .Rana sierramadrensis (upper left, MSU 12660, 51.3 mm S VL, 1 1 km. S Chicahuaxtla,
Oaxaca), Rana sinaloae (upper right, MSU 12661 , 75.8 mm S VL, 22 km. WSW Ameca, Jalisco),
and Rana maculata (bottom, AMNH, 88 mm S VL, Finca El Rosario Vista Hermosa, ca. 7 km N
Escuintla Guatemala, from kodachrome by Richard G. Zweifel), for comparison in pattern.

1978

A Systematic Review of the Mexican Frog

11

(but usually only two), no marginal teeth, more extensively pigmented dorsal tail fins,
and in having the lateral margins of the oral disc emarginate. The larvae of R. pus-
tulosa usually have some dark marks and blotches on the dorsal tail fins and mus-
culature (mostly clear in R. sierramadrensis ) and at least distally on the ventral fins
(virtually lacking in R. sierramadrensis)', in some larvae of R. pustulosa with relatively
clear dorsal fins small punctate melanophores are more extensive than in R. sierra-
madrensis. The oral disc is relatively smaller in R. pustulosa (about two-thirds) than
in R. sierramadrensis, and the lateral margins are infolded in larvae of/?. pustulosa
(see comparison in figure 3 A, B). Less reliable is the number of upper tooth rows in
distinguishing the two species, which is usually five (maximal at six) in larvae of R.
pustulosa, and which is probably six but certainly seven in larvae of R. sierramad-
rensis. The larvae of R. sinaloae resemble those of R. sierramadrensis only in having
a mostly clear dorsal tail fin; the larvae of R. sinaloae otherwise are not different from
those of R. pustulosa and are distinguished from larvae of R. sierramadrensis by the
same features that differentiate those two species.

The larvae of R. sierramadrensis are easily distinguished from larvae of both R.
maculata ( macroglossa ) and R. palmipes (comparative data in Volpe and Harvey,
1958) in having only three lower tooth rows, and in the mostly clear tail fins. The
larvae of R. sierramadrensis may have more highly arched dorsal fins than larvae of
R. maculata. The larvae of/?. sierramadrensis and/?, maculata otherwise share sev-
eral features that distinguish them from larvae of R. palmipes. The larvae of both
species have a maximum of six or seven upper tooth rows (four or maximum of five
in R. palmipes ), have a narrow medial A-2 gap (wider in/?, palmipes ), have marginal
teeth (lacking in/?, palmipes ), and have a suctorial oral disc that lacks infolded lateral
margins (infolded in/?, palmipes ).

Relationships

Rana sierramadrensis is currently placed in the Rana palmipes species group
(Zweifel 1954). So far as known the white, spinous excrescences in males of/?, sier-
ramadrensis are unique. One seemingly trenchant feature is shared with the member
species of the/?, tarahumarae group — three lower rows of teeth in larvae (the number
may be variable in /?. sierramadrensis, but is invariably three in R. tarahumarae
group). All other member species of the /?. palmipes group have larvae with four
lower tooth rows (/?. palmipes, R. maculata, R. vibicaria, R. warschewitschii)', Lee
(1976) referred some peculiar tadpoles from Belize with five lower tooth rows to/?.
maculata .

Variation in the two species/?, palmipes and/?, maculata (nearest geographically
to/?, sierramadrensis) has not been investigated in detail. In addition to other features
distinguishing the two species, /?. palmipes is especially distinctive from /?. sierra-
madrensis in males having enlarged, glandular thumb pads; males of /?. palmipes
based on statements by Boulenger (1920:417), Taylor (1952:908), and Zweifel
(1967:54) seem to have internal slits but lack external vocal sacs. Rana maculata
differs from /?. sierramadrensis in having well-developed vocal sac slits, but resem-
bles/?. sierramadrensis in having the thumbs “not larger than in females” (Smith
1959:216); presumably males of/?, maculata lack the white, spinous excrescences of

12

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No. 300

R. sierramadrensis. Rana sierramadrensis seems to be most closely allied to, and the
trans-isthmian counterpart of, R. maculata. Aside from the above-mentioned differ-
ence and those that readily distinguish the larvae (tail pigmentation and number of
lower tooth rows), both species share general features of pattern (Fig. 5), occupy
upland habitats, and have larvae with stream-adapted modifications. In pattern, R.
maculata differs from R. sierramadrensis in having wider dark bars on the femora,
in usually lacking a continous black, ventrolateral band, and (at least in some specimens)
in having a green dorsum (Kodachrome slide of R. G. Zweifel, Fig. 5).

RESUMEN

La variacion de Rana sierramadrensis se discute. Los renacuajos son descritos
por primera vez. Una caracterfstica secundaria sexual no conocida es la presencia de
tuberculos espinosos blancos en los no engrandecidos pulgares, pecho, y mandfbulas
inferiores de los machos. Rana sierramadrensis se retiene en el grupo/? . palmipes. Rana
sinaloae se traslada al grupo R. tarahumarae, qual se propone para inclusion de las
especiesR. tarahumarae, R. pustulosa, R. sinaloae, R. pueblae, y R. johni, Rana
sierramadrensis ocurre solamente en la Sierra Madre del Sur en los estados Mexicanos
de Guerrero y Oaxaca, y parece ser relacionada aR. maculata.

LITERATURE CITED

Altig, R. 1970. A key to the tadpoles of the continental United States and Canada. Herpeto-
logica 26:180-207.

Boulenger, G. A. 1920. A monograph of the American frogs of the genus Rana. Proc. Amer.
Acad. Arts Sci. 55(9):41 3-480.

Case, S. M. 1976. Biochemical systematics of Rana in western North America. Herp. Review
7(2): 76 (Abstract).

Cochran, D. M. 1961 . Type specimens of reptiles and amphibians in the United States National
Museum. Bull. U.S. Nat. Mus. No. 220, xv -f- 291 pp.

Davis, W. B. and J. R. Dixon. 1959. Snakes of the Chilpancingo region, Mexico. Proc. Biol.
Soc. Washington 72:79-92.

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on
identification. Herpetologica 16:183-190.

Lee, J. C. 1976. Rana maculata Brocchi, an addition to the herpetofauna of Belize. Herpeto-
logica 32(2):21 1-214.

Rugh, R. 1962. Experimental embryology. Ed. 3, Burgess Publ. Co., Minneapolis, Minnesota,
pp. ix + 501.

Smith, H. M. 1959. Herpetozoa from Guatemala. I. Herpetologica 15:210-216.

, D. A. Langebartel, andK. L. Williams. 1964. Herpetological type-specimens in the

University of Dlinois Museum of Natural History. Illinois Biol. Monogr. No. 32, Univ.
Illinois Press, 80 pp.

Taylor, E. H. 1939. New species of Mexican tailless Amphibia. Univ. Kansas Sci. Bull. (1938)
25(17):385-405.

1944. Present location of certain herpetological and other types (sic) specimens. Univ.

Kansas Sci. Bull. 30(1 1): 1 17— 187.

1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull. 35(5):577-942.

1978

A Systematic Review of the Mexican Frog

13

Volpe, E. P. and S. M. Harvey. 1958. Hybridization and larval development in Rana palmipes
Spix. Copeia 1958(3): 197-207.

Zweifel, R. G. 1954. A new frog of the genus Rana from western Mexico with a key to the
Mexican species of the genus. Bull. Southern California Acad. Sci. 53(3):131 — 141 .

1955. Ecology, distribution, and systematics of frogs of th eRana boylei group. Univ.

California Publ. Zool. 54(4):207-292.

1967. Systematic status of the Central American frog, Rana miadis. Herpetologica

23( 1):54— 56.

Accepted for publication March 16, 1977.

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