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Volume 85 Number 1

BCAS-A85(1) 1-64 (1986) APRIL 1986

Southern California Academy of Sciences

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© Southern California Academy of Sciences, 1986

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Date of this issue 18 April 1986

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 1-15
© Southern California Academy of Sciences, 1986

Ecology of the Desert Kit Fox, Vulpes macrotis arsipus,
in the Mojave Desert of Southern California

Thomas P. O’Farrell! and Larry Gilbertson?

‘EG&G Energy Measurements, Inc., 130 Robin Hill Road,
Goleta, California 93117
(mailing address: 611 Avenue H, Boulder City, Nevada 89005), and
2?Box 591, McGill, Nevada 89318

A field study using live-trapping and radio-telemetry was conducted to learn
whether desert kit foxes inhabiting the Rand Open Area in eastern Kern County,
California, were being adversely affected by human activities such as use of ORV’s,
as compared with foxes living in the undisturbed Desert Tortoise Research Natural
Area. Individuals were more readily trapped in the Rand Area, and fox populations
there were more dense, had a higher rate of reproduction, greater survival, and
different sources of mortality as compared with the control population in the
Tortoise Area. Except for two foxes shot in the Rand Area, there were no negative
effects observed in the ORV study site that could be causally linked to human
activities.

The kit fox, Vulpes macrotis, is a widely distributed predator in the arid habitats
of western North America. Of the eight subspecies named since 1888, V. m.
macrotis, which was first trapped in Riverside County, California, was extinct by
1910 (Grinnell et al. 1937). In the San Joaquin Valley of California V. m. mutica
lost a significant portion of its original habitat through increased agricultural,
mineral, urban, and industrial developments (Laughrin 1970; Morrell 1972, 1975).
This justified its receiving Federal protection as an endangered species following
passage of the Endangered Species Protection Act in 1966 (Federal Register, 32:
4001), and its classification as a rare species by the California Department of Fish
and Game (State of California 1980).

The status of other subspecies of V. macrotis was poorly known. This concerned
the Desert Planning Staff of the U.S. Bureau of Land Management (BLM) who
was developing comprehensive land-use plans for the deserts of southern Cali-
fornia (Anonymous 1980). Without appropriate baseline information it would be
difficult, if not impossible, to effectively manage essential habitats necessary for
the well-being of the desert kit fox, V. m. arsipus, and other potentially vulnerable
species of wildlife.

It was known that the kit fox is a relatively unwary animal (Hall 1946; Egoscue
1956, 1962; Laughrin 1970). Highway fatalities are a significant source of reported
mortality (Egoscue 1962; Morrell 1972), and kit fox are also vulnerable to being
shot at night by predator hunters (Laughrin 1970; Morrell 1972). The use of
poisoned grains to control rodents in California poses a potential serious threat
because kit fox may become secondary, nontarget victims if they consume either
poisoned rodents or bait (Swick 1973; Schitosky 1975). It has also been suggested

1

2 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

that indiscriminate killing and disturbance of kit fox habitats and prey by off-
road vehicle (ORV) enthusiasts has a serious, negative effect on the species (Laugh-
rin 1970).

The major goal of this project was to provide BLM with information on the
ecological life history requirements of the desert kit fox in the Mojave Desert of
southern California that would be useful in their planning document. Specific
goals were to: examine habitat preferences, locate and characterize denning sites,
determine denning range and movement patterns, and document sources of mor-
tality in undisturbed desert habitats and those being affected by human activities
such as ORV use.

Methods
Study Areas

Two study areas were selected in eastern Kern County, California. The first was
located on the Desert Tortoise Research Natural Area, approximately 3 km north-
east of California City (Fig. 1). This area was partially fenced and access and
grazing were restricted. Although the site had a past history of disturbances by
sheep grazing and wildfires, it served as a “control” area that was relatively free
from contemporary human disturbances.

The second study area was located in the BLM’s Rand Open Area which was
adjacent to the northeastern boundary of the Tortoise Area. “Indiscriminate”’
ORV uses were permitted there. It was chronically and heavily grazed by sheep,
and in some areas it adjoined agricultural developments. The Rand Area served
as the treatment plot where kit fox populations were subjected to periodically
intensive human interference.

Both areas had similar elevational gradients (580-900 m). Soils below 610 m
were light-textured, while those at higher elevations had coarser texture due to
higher concentrations of granitic fragments. Near the ridge lines, soils were shal-
lower and contained larger rock fragments and outcrops.

The dominant vegetation association would have been Larrea-Ambrosia with
an understory of native forbs and grasses. However, years of grazing have reduced
the sites to a Larrea-Schismus-Erodium disclimax. The few native winter annuals
observed in 1978 were under the canopies of either unpalatable or thorny shrubs.
There were few native plants germinating between shrubs. Ambrosia dumosa
contributed far less to the plant cover than expected, since most individuals had
either been eaten or trampled.

Live-trapping

Kit fox were captured using two types of collapsible, wire mesh live-traps:
double-door traps measuring 38 < 38 x 107 cm, or single-door traps measuring
30 x 30 X 76 cm. Traps were usually placed at or near kit fox dens, but on a few
occasions they were operated near intersections of dirt roads or in washes away
from dens. Bait such as sardines in cheesecloth bags, chicken bones, road-killed
birds or rodents, or pieces of freshly killed jackrabbits (Lepus californicus), was
wired to the back door or top of traps.

Captured foxes were handled as quickly and humanely as possible to minimize
trauma. A heavy pillowcase was used to distract the fox so that it could be quickly

ECOLOGY OF THE DESERT KIT FOX 3

4

INTENSIVE
STUDY AREA

OREGON
Miles

2 3h 5 6

6 Ss IO

Kilometers

Fig. 1. Location of kit fox study sites in Rand Open Area and Desert Tortoise Research Natural
Area, Kern County, California. Dens DT-23 and RA-33 are the first capture and final recovery sites
for a dispersing female.

grasped by the nape and withdrawn from the trap. When held by the nape most
foxes became submissive and ceased struggling. The animal was placed on the
ground with the pillowcase covering its eyes, and its rump and hind legs were
firmly held between the knees and legs of the kneeling investigator.

A numbered, Monel metal ear tag supplied by the California Department of
Fish and Game was secured to each fox’s ear to provide long-term identification
of individuals. The right ear was measured from the notch, and observations of
the animal’s sex, reproductive status, and overall condition were recorded. Fleas
and other ectoparasites were observed to determine relative numbers before sam-

4 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ples were taken and preserved in alcohol. Animals were weighed on a portable
scale and released at the point of capture.

Radio-telemetry

Selected animals were fitted with lightweight (100 g) radio transmitters (Telon-
ics, Inc., Mesa, AZ) operated at individual frequencies between 150.2 and 151.1
MHz. Each was equipped with an integral, flexible, whip antenna, and was secured
to a collar made of a urethane-coated nylon webbing.

Animals were tracked using a multichannel receiver equipped with either an
omni-directional antenna, or a multielement, hand-held yagi antenna, and stan-
dard radio-telemetry techniques. The directional yagi antenna was also attached
to the strut of a Super Cub during aerial surveys to locate animals whose signals
were undetectable during routine ground reconnaissance.

Den Descriptions

Kit fox dens were described by observing and recording: number of entrances;
elevation; soil type; major vegetation associations; distance to nearest mappable
road; and the presence of fresh fox tracks, scats, and prey remains. The den was
judged to be either active or inactive based on the freshness of sign. A unique
code number was assigned and the location of the den was plotted on a topographic
map using compass bearings on prominent land features.

Occupancy of dens, especially by mated pairs, was determined by locating as
many radio-collared foxes in their dens as possible each day.

Necropsies

Dead foxes were located as quickly as possible so that information could be
gathered on probable cause of death. External measurements, including total
length, tail, right hind foot, and right ear from notch, and weight were taken for
all animals that were not mummified. External conditions were noted, especially
the presence of injuries, blood, ectoparasites, feces, and obvious anomalies. A
mid-ventral incision from the larynx to the urogenital openings was made on
animals that were not badly decomposed. Each organ and various tissues were
observed for gross lesions and evidence of decomposition. The gastrointestinal
tract was checked for food items and endoparasites. General condition of repro-
ductive organs was observed.

Results
Live-trapping

Between November 1977 and January 1979, 13 foxes (six males, seven females)
were captured 20 times in the Rand Area, for an overall trapping success of 34%.
Eight foxes were captured only once, three foxes were recaptured once each, while
two foxes were recaptured twice each. Although trapping effort was greater in the
Tortoise Area (149 versus 59 trap/nights), trapping success was only 9%. Thirteen
individuals (eight males, five females) were captured; none were ever retrapped.

Radio transmitter collars were fitted to six foxes (three males and three females)
in the Rand Area, and to all 13 foxes live-trapped in the Tortoise Area. No data
were gathered that indicated that the collars were too heavy or interfered with
normal activities of the foxes.

ECOLOGY OF THE DESERT KIT FOX 5

Table 1. Number of entrances to kit fox dens observed in the Rand and Desert Tortoise areas,
Kern County, California, 1977-1979.

Number of Rand Area dens Tortoise Area dens
entrances (treatment) (control)
1 5 8
2 7 10
3 7 7
4 1 3
5 2 4
6 2
7 3 1
8 yD) 1
9 1
12 1
18 1
22 a a
Total 33 34
Mean (+SE) 5.0 + 0.8* 2.8 + 0.3
Median 3 2

* Statistically significant difference between means (P < 0.05).

Complete records of trapping information and radio telemetry relocation data
were published in Appendix | of the study report (O’Farrell and Gilbertson 1979).

Dens

A total of 33 dens were found in the Rand Area. Only seven were found by
observers hiking or driving through the area; most were found by tracking radio-
collared foxes to new dens (157 radio-fixes). Of 34 dens found in the Tortoise
Area, 14 were found by observation, while tracking radio-collared animals re-
vealed 20 new dens (131 radio-fixes).

The average number of entrances to dens was 5.0 + 0.8 (mean + standard
error) and 2.8 + 0.3 for the Rand and Tortoise areas respectively (Table 1). The
significantly greater (P < 0.05) number of entrances to Rand Area dens was also
indicated by the fact that 33% of the dens there had more than five entrances,
and that the largest dens observed (>10 entrances) were all found there. Only
two dens located in the Tortoise Area had more than five entrances.

Mean elevation of dens in the Rand Area was 667 + 7 m (range 585-770 m),
which was significantly lower (P < 0.05) than the average elevation of 733 + 8
m (range 675-830 m) for the Tortoise Area dens. Almost all of the dens were
located on gradual, west- to northwest-facing slopes dominated by a Larrea-
Schismus-Erodium disclimax. Soils were characteristically deep, light-textured,
and virtually stoneless to depths of 1.5 m. There was no observable caliche or
hardpan layer, and dens were usually placed in well drained sites.

The average distance between dens and the nearest dirt road or jeep trail was
207 + 42 m and 163 + 38 m for the Rand and Tortoise areas, respectively (Table
2). Only four dens, two in each area, were located more than 500 m from a road.
More than half of the dens on the Tortoise Area were less than 50 m from a road
or trail.

Three atypical dens were found. A mated pair in the Rand Area occupied a

6 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 2. Distance from kit fox dens to nearest dirt road or trail in Rand and Desert Tortoise areas,
Kern County, California, 1977-1979.

Distance to
road (m) Rand Area dens Tortoise Area dens
0-25 2 12
26-50 4 6
51-75 3
76-100 6 2,
101-150 3 3
151-200 6 4
201-300 4 1
301-400 1 1
401-500 2 D
501-600 1
601-700
701-800 1 D
1200 1
Mean (+SE) 207 + 42 163 + 38
Median 150 43.5

single-entrance den placed between horizontal rock layers on the north-facing
slope of a small hill at an elevation of 690 m. Another den was located in a culvert
under a well-used access road to a recreational vehicle rendevous and camping
site in the Rand Mountains. A third den was found next to a steel well casing on
the Desert Tortoise Area. The soil adjacent to the casing was eroded in places to
a depth of 1.5 m. The fox gained access to a horizontal den opening after dropping
down along the casing.

All but four of the 33 dens located in the Rand Area were occupied by radio-
collared foxes at some time. Of the occupied dens, 18 were used by single foxes,
10 were used by two foxes (nine by mated pairs, one by two lone foxes), and one
was used by three collared foxes (one pair and one lone fox). In the Tortoise Area
14 of 34 dens were never used by foxes carrying radio collars. Of the 20 dens that
were used, 14 sheltered only one collared fox each, four were occupied by two
foxes (two mated pairs in two dens; two unpaired foxes in each of two dens), and
two were occupied by three different foxes (one den used by mated pair; one den
used by three lone foxes).

Six foxes in the Rand Area used an average of 7 + 1.3 dens each with a range
of six to nine (Fig. 2). Approximately 42% of the dens were shared by mated
pairs. In spite of their proximity, mated pairs appeared to use a series of dens
exclusively from their neighbors (Fig. 2). Only one female used more dens alone
than with her mate, but most of these were occupied after he was killed.

Three large pupping dens (dens known to have been occupied by mated pairs
and their puppies) were located in the Rand Area. They were an average of 2 km
apart.

The sizes of dens occupied by mated pairs in the Rand Area, indicated by the
number of entrances, differed during and after the breeding season. From January
to the end of May, the last time puppies were observed with the pairs, the average

ECOLOGY OF THE DESERT KIT FOX 7

PA
272 FEMALE
339 MALE

274 FEMALE
341 MALE

340 FEMALE
342 MALE

F—1 KILOMETER—4

1 MtLE—————}

Fig. 2. Locations of dens in Rand Area occupied by mated pairs of kit fox either together (closed
circles) or alone (solid stars). Solid line encloses approximate denning ranges. Hatched area indicates
only known place where ranges of two pairs may have overlapped. Numbers are den identification
codes. Contours are in feet, and dashed lines indicate dirt roads.

number of entrances to shared dens was 8.5 (range 5-18). This was significantly
greater (P < 0.05) than the average of 3.4 entrances (range 1-8) observed for
shared dens occupied between June and December.

Between one and three dens were occupied by the 13 radio-collared foxes in
the Tortoise Area, for an average of 2.1 + 0.8 dens occupied per animal. Only
six of the foxes were known to share their dens with other radio-collared foxes;
the majority of foxes lived alone or with uncollared mates.

When den sites for six foxes in the Rand Area and for 10 foxes in the Tortoise
Area were mapped, the greatest distance between dens used by the individuals
averaged 1.6 + 0.4 km. If the average longest distance were considered to be the
estimated diameter of a circular home range, the mean denning range would
occupy 200 ha. The denning range was defined as the area enclosing the den sites
occupied, usually exclusively, by individual foxes.

Complete descriptions of dens were published in Tables 3 and 4 of the project
report (O’Farrell and Gilbertson 1979).

8 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Dispersal

At least two animals moved distances that indicated they were dispersing rather
than moving within established home ranges. A male was first caught 2 April
1978 along the west-central border of the Tortoise Area. Over the next eight
months he was located aboveground several times, and in a series of dens between
the Tortoise Area and California City. The greatest distance between sightings
was 4.8 km. A female was found in a den at the base of the El Paso Mountains
approximately 32 km northeast of her initial capture point in the Tortoise Area
a month earlier (Fig. 1).

Pairing and Reproduction

Three pairs in the Rand Area and one pair in the Tortoise Area were presumed
to be mated pairs. The pair in the Tortoise Area was together during March, but
both animals died shortly thereafter.

One pair in the Rand Area was together for at least three months, April through
June. Puppies were observed in their den in late May. They apparently separated
in mid-June, and the male was killed in late July or early August.

Another pair was together throughout the study. They were in common dens
from April 1978 through January 1979. Puppies were observed in their dens in
late May also. Neither was ever observed in dens with another radio-collared fox.

A third pair was observed to be in the same dens from April through at least
late November 1978. At that time they either dispersed far enough to be out of
radio range, or their radios both quit 3—4 months prematurely, and they were
either untrappable or dead at the termination of the field studies.

Adult males had palpable scrotal testes throughout the year. During January
the penises of reproductively active males were obviously engorged with blood.
An exception was an unpaired male whose penis was noticeably smaller, un-
swollen, and flesh-colored, suggesting that he was not breeding. His weight (2025
g) was comparable with that of breeding males at the time.

Adult females were also reproductively active in January. They had swollen,
moist vulvas, and some had already started to make weight gains suggesting that
they may have been pregnant. Lactating females with swollen mammae and mat-
ted hair around their conspicuous teats were observed in March and April. Puppies
were first observed aboveground near their natal dens in late May. None were
observed with their parents after that.

Longevity

Longevity was defined as the amount of time foxes were known to be alive. It
was calculated from the date of first capture to either the date of the last capture
or the date of the last location obtained using radio-telemetry. In the Rand Area
the six foxes caught prior to the last trapping session in January 1979, had an
average longevity of 10.2 + 0.8 months with a range of between 8-14. Four of
the animals were still alive when the study was terminated. Another whose radio
presumably malfunctioned was probably alive as well.

Thirteen foxes in the Tortoise Area had an average longevity of just 2.4 + 0.7
months. Seven of the animals were known to be alive for 30 days or less, and

ECOLOGY OF THE DESERT KIT FOX 9

Table 3. Observed sources of mortality in kit fox, Kern County, California, 1977-1979.

Area Number Probable cause of death

Rand Area (treatment) 2 Shot
Tortoise Area (control)

Found underground 2 Suffocated; den cave-ins
3 Emaciated; starved and/or diseased (?)
3 Unknown
Found aboveground 1 Severe trauma to skull, thorax; predator
1 Severe trauma to thorax; predator,
probably coyote
Both areas
(tagged Tortoise Area found under- 1 Emaciated, broken hind leg; multiple
ground Rand Area) probable causes

only one animal lived as long as nine months. Average longevity in the Tortoise
Area was significantly shorter (P < 0.05) compared with average longevity in the
Rand Area. None of the animals was known to be alive at the termination of the
study.

Mortality

The radio-collars permitted finding 12 dead foxes that were still in a suitable
condition to determine probable cause of death, and a dead, untagged animal was
found incidentally (Table 3). Two males were found shot to death in the Rand
Area. One was on the surface about 150 m from its most recently occupied den.
The other, an untagged animal, was found in the entrance to a den formerly used
by a mated pair where he had been stuffed by a hunter who had also cut off his
tail.

Of the 10 animals found dead in the Tortoise Area, eight were found under-
ground in their dens: three died of unknown causes; three had no food in their
stomachs, no subcutaneous or mesenteric fat, and appeared to have starved; two
animals were found in collapsed tunnels and probably suffocated in cave-ins.

Two animals were found on the surface. One had its skull crushed and other
signs of traumatic death due to a predator; the other had been killed so recently
that its body fluids were still near body temperature and the carcass was shipped
to the Wildlife Investigations Laboratory, Wildlife Management Branch, Cali-
fornia Department of Fish and Game. The examining veterinarian’s diagnosis
was, “Thoracic trauma, probably dog or coyote. Emaciation ... .”

The female that dispersed 32 km was also found dead underground. She was
also emaciated and had a compound fracture of the right femur. The new den
was between two primary roads and she may have been hit by a vehicle. This
probably did not kill her directly, however, since there were no signs of internal
trauma or hemorrhaging that usually accompany road-kills.

Measurements

Standard measurements (in mm) of foxes found dead in the Tortoise Area were:
(three males) total length, 797 + 9; tail, 320 + 10; right hind foot, 127 + 1; right

10 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ear from notch, 86 + 2; (three females) total length, 772 + 12; tail, 302 + 5;
hind foot, 125 + 2; ear from notch, 80 + 1. Average live weights were: males
(15), 2034 + 78 g; females (13), 1936 + 64 g.

Fleas

A total of 299 fleas representing five species was identified in 26 collections.
The most common were the so-called human fleas, Pulex irritans (107 males, 132
females), and the poultry or sticktight fleas, Echidnophaga gallinacea (26 males,
27 females). One male Orchopeas leucopus, one female O. sexdentatus, and two
male and three female Cediopsylla inequalis were also collected.

Abnormalities

Four abnormalities, none of which resulted from the trapping or tagging op-
eration, were observed: a broken left canine; a hardened scar on a foreleg; a reddish,
glazed eye that may have been injured; and a ruptured cornea that also may have
been due to a recent injury.

Discussion

Results of the live-trapping efforts were the first indication that populations of
foxes in the Rand and Tortoise areas were probably not comparable. Foxes were
readily trapped in the Rand area, as shown by the high overall trapping success.
Several animals were recaptured, two of them twice each after their initial captures.
In spite of increased efforts in the Tortoise Area, the overall trapping success was
only 9% and none of the animals were recaptured. Trapping data suggested that
the fox population in the Tortoise Area was less dense than in the Rand Area.

Kit fox dens in both areas occupied sites having similar soils, exposure, ele-
vation, and vegetation associations. Soils were always loose-textured and free of
rocks or stones. Most authors have associated the kit fox with loose-textured soils
(Burt 1934; Hall 1946; Egoscue 1962; Laughrin 1970; Morrell 1972). Morrell
(1972) speculated that it might be difficult for kit fox to dig in soils with a hardpan
unless other animals, such as badgers, penetrated the impervious layer.

Few of the dens were found in the sandy soils at elevations below 670 m, and
none were found at elevations above 820 m where the soils were generally shallow
and rocky. The significant difference between mean elevation of den sites on the
two study areas probably reflected the fact that optimal textured soils reached
lower elevations in the Rand Area compared with the Tortoise Area.

It was anticipated that kit fox would select areas remote from human habitations
and disturbances for den sites. Surprisingly, this was not the case, as most dens
were within 150 to 200 m of roads and jeep trails, and few were found to be more
than 300 m away. It is possible that the human activities conducted on the roads
did not disturb the foxes significantly. It is also possible that the dirt roads and
trails provided foxes with additional paths along which the foxes had increased
visibility for hunting and for remaining vigilant against predators.

Kit fox dens in the Rand Area had a significantly greater number of entrances
than dens observed in the Tortoise Area. If the premise is accepted that older,
natal, or pupping dens have more entrances than younger, daily-use shelter dens,
then the Rand Area provided a more optimal habitat than the Tortoise Area as
regards both potential breeding sites and dens to rear puppies. This inference is

ECOLOGY OF THE DESERT KIT FOX 11

further supported by the lack of evidence of successful breeding in the Tortoise
Area, and absence of puppies observed there.

In Utah, kit fox dens had between two and seven entrances, although one den
had 24 (Egoscue 1962). Dens of V. m. mutica in western Kern County had one
to 10 entrances, although most had just two (Morrell 1972). When the average
number of entrances to dens was compared between subspecies, dens of V. m.
arsipus in eastern Kern County were more similar to dens of V. m. nevadensis in
Utah than they were to dens of V. m. mutica in nearby western Kern County.

Although kit fox prefer to construct underground dens, they were observed
denning adjacent to a well casing and in a culvert. Egoscue (1956) also reported
that kit fox occasionally used culverts as dens. The den located between rock
layers on a low hill was the most unusual since the single entrance appeared to
be in solid rock.

In the Rand Area, 88% of the dens were used by radio-collared foxes. Most of
the occupied dens were used either by lone foxes or mated pairs. In the Tortoise
Area only 59% of the known dens were ever occupied by collared foxes. Egoscue
(1962) also reported that study areas in Utah contained a mixture of occupied
and deteriorating dens.

Individual foxes in the Rand Area used more dens than animals on the Tortoise
Area. The difference in the average number of dens occupied was probably due
to the short duration that radio-collared foxes remained or lived in the Tortoise
Area. If more foxes had lived longer there, they may have used as many dens as
their neighbors in the Rand Area.

That mated foxes occupied larger dens during the pupping season than during
the remainder of the year confirmed Morrell’s (1972) finding that San Joaquin
kit fox occupied smaller dens during the three months following the breeding
season. The importance of the larger natal or pupping dens for successful breeding
and survival of the species is only now being appreciated (Egoscue 1962; Morrell
1972).

The three natal or pupping dens occupied by neighboring mated pairs were
about 2 km apart. Those in Utah were at least 3.2 km apart (Egoscue 1956, 1962).
Spacing of natal or pupping dens may be important in the territorial requirements
of the species, especially since data on den-use patterns supported earlier obser-
vations that mated pairs used certain sets of dens exclusively (Morrell 1972). This
suggests a territorial allocation of available habitat. Egoscue (1975) believed that
the maximum carrying capacity of an area for foxes was primarily related to their
territorial requirements.

During this study the estimated denning range was 200 ha. This is comparable
with Morrell’s (1972) estimate of 260 to 520 ha for V. m. mutica. Although Morrell
called his estimate a home range, an examination of original data (Morrell 1971)
showed that he drew boundaries around occupied dens (a denning range), but had
no additional information on movements of foxes away from den sites to hunt
or maintain territorial boundaries (home range).

The longest dispersal movement recorded was identical in length with two 32-
km dispersal movements recorded by Egoscue (1962) in Utah. He showed that
puppies did not establish residence in the area of their birth. The dispersal move-
ments recorded in Utah and California may have represented dispersal of juveniles
from their natal sites to new ranges.

12 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Timing of reproduction in kit fox in the Rand Area was similar to that described
for other subspecies (Egoscue 1956, 1962; Morrell 1972). Adult males had palpable
scrotal testes throughout the year. The most conspicuous change in their genitalia
occurred in January when their penises were obviously enlarged, red in color, and
often appeared to be extended from their sheaths. Females were in estrus in
December and January. Evidence based on weight gains of vixens in January, and
observations of lactation in March suggested that puppies were born in February
or March.

In other studies breeding took place between December and March (Egoscue
1956), and an average of four to five puppies were born between February and
March (Egoscue 1956, 1962; Morrell 1972). Puppies did not appear aboveground
until they were at least one month old (Morrell 1972).

Kit fox are thought to be essentially monogamous, and may mate for life
(Grinnell et al. 1937; Egoscue 1962). Information gathered during this study, as
well as the results of previous studies (Egoscue 1962; Morrell 1972) indicated that
kit fox employ a flexible strategy as regards reproductive pairing. Some pairs may
remain mated for more than one breeding season, while others may change part-
ners frequently. Egoscue (1962) reported that most mated pairs remained bonded
but he did observe three cases of polygamy. Morrell (1972) observed that only
one of seven mated pairs remained bonded the next breeding season.

Information on longevity provided further evidence that the populations of kit
foxes on the two areas were not comparable. In the Rand Area most of the adults
tagged at the onset of the study were still alive at its termination. In contrast,
most of the foxes in the Tortoise Area were known to be alive for one month or
less, and none was alive when the study was terminated. The relatively high
turnover of adults in the Tortoise Area, compared with stability of adults in the
Rand Area, suggested that the Tortoise Area was not a suitable habitat for the
species between 1977 and 1979. The factors responsible for differences in longevity
between the two areas were not obvious, but information on sources of mortality
suggested tentative answers.

In the Rand Area only two dead foxes were retrieved; both had been shot. The
remaining animals were known to be, or were presumably, still alive at the end
of the study.

The high proportion of foxes in the Tortoise Area that showed evidence of
starvation, plus the absence of similar data in the Rand Area, suggested that food
supplies were more dependable in the latter area. Morrell (1972) speculated that
starvation was a factor limiting kit fox populations, and he further suggested that
the total population may decline in years of low rodent populations. Red fox (V.
vulpes) also starve during food shortages rather than shifting their territories to
sites with more numerous prey (Scott and Klimstra 1955).

Although starvation was the proximate cause of death in several of the kit fox
in the Tortoise Area, the ultimate cause of death may have been a contagious
disease such as canine distemper. One of the clinical symptoms of this and other
viral and bacterial diseases is anorexia and a gradual loss of activity (Armstrong
and Anthony 1942; Huff et al. 1974). The loss of almost every animal in the
Tortoise Area in a brief period of time, the fact that they died underground, the
absence of food in their gastrointestinal tracts, their emaciated overall condition,
and the presence of diarrhea and conjunctival secretions provided circumstantial

ECOLOGY OF THE DESERT KIT FOX 13

evidence that a disease may have been the ultimate cause of death. Unfortunately,
the animals were not recovered quickly enough to provide tissue samples for
histopathological diagnoses.

In the Tortoise Area two adult foxes were dug from their dens after they ap-
parently suffocated in cave-ins. Although the role of accidents as a source of
mortality in wild animals is seldom considered, it certainly appeared that the
headwalls of both dens collapsed, pinning the animals in chambers. The remaining
tunnels appeared to be clear and unaffected by the cave-ins. Natural ground
movements, such as earthquakes, plus man-made shocks, such as heavy ORV’s
bouncing above a den, may play a role in causing cave-ins and mortality of kit
foxes.

Two foxes died due to trauma caused by a predator. Teeth on a reference skull
of a coyote fit the width of the puncture wound found between ribs of the crushed
thoracic cavity of one animal, and the consulting pathologist concluded that at
least one of the foxes died of a coyote or dog bite. Coyotes probably killed both
animals because there were no feral or domestic dogs observed in the vicinity
when the animals were found. Neither animal, however, was eaten by its killer.

Coyotes are considered to be potential predators of kit fox (Goldman 1930;
Knapp 1978; Young and Jackson 1951), and Seton (1937) offered the thesis that
kit fox used dens to avoid confrontations with coyotes. Egoscue (1962) was unable
to provide conclusive evidence that coyotes were, or were not, predators of kit
fox. He did, however, relate that a coyote was observed digging into a kit fox den
that was found to contain a small puppy. Another coyote was observed stalking
a fox family that was aboveground near their den. However, kit fox remains were
found in only two of approximately 8357 coyote stomachs examined (Sperry
1941), which suggests that if they kill kit foxes, coyotes seldom if ever consume
them.

Animals in both areas were comparable as regards standard measurements and
average weights. Standard measurements compared favorably with reported val-
ues (Grinnell et al. 1937; Hall 1946), but they failed to confirm that standard
measurements of females are 2—7% smaller than those of males (Grinnell et al.
1937). It was also reported that male V. m. mutica were 15% heavier than females
(Grinnell et al. 1937), and that male V. m. nevadensis were 8% heavier than
nonpregnant females (Egoscue 1962). There was no significant sexual dimorphism
in weights of V. m. arsipus.

Fleas were the most conspicuous ectoparasites observed on kit fox. They were
found throughout the year, but appeared to be most dense between December
and April. They also seemed to be more dense on females than on males, especially
during the pupping season. The human flea was the most common and numerous
species. It was also found to be the most common flea on kit foxes in Utah
(Egoscue 1962).

The sticktight flea was found on 35% of the foxes sampled. It has been found
on kit foxes in Arizona (Turkowski 1974), California (Morrell 1972), and Nevada
(Beck and Allred 1966), but never on kit foxes sampled in Utah (Egoscue 1962).
It is more commonly found on birds.

The few other fleas found were probably prey fleas that transferred to foxes
when their preferred host was killed or consumed. Orchopeas leucopus is most
commonly found on deer mice (Peromyscus maniculatus), and harvest mice

14 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

(Reithrodontomys megalotis); O. sexdentatus is usually found on pack rats (Neot-
oma spp.); and Cediopsylla inequalis is most often found on lagomorphs, especially
cottontails (Sy/vilagus spp.).

Except for the shooting of two foxes in the Rand Area, there were no significant,
immediate negative effects observed in the ecological life history parameters under
study that could be causally linked to human activities such as ORV use. Kit fox
populations there were more dense, appeared to have a higher rate of reproduction
and greater longevity, and did not appear to suffer from either starvation or disease,
as compared with the fox population in the relatively undisturbed Tortoise Area.

The choice of the Tortoise Area population as a “‘control’’ may have been
inappropriate if kit foxes there were being subjected to severe natural stresses,
such as low food supplies or infectious diseases, that were not concurrently af-
fecting the Rand population.

The question of how ORV’s and other human activities influence kit foxes will
not be satisfactorily answered until adequate paired populations are found, and
suitable field experiments are designed and executed to apply measured ORV
stresses to kit foxes in their dens.

In the meantime, large natal or pupping dens should be protected from adverse
human activities or destruction. Results of this and other published studies suggest
that natal or pupping dens, used exclusively by mated pairs, may be an important
element in the critical habitat of the species.

Acknowledgments

The field study was sponsored by the U.S. Bureau of Land Management, Desert
Planning Staff, through contracts CA-060-CT8-55 and CA-060-CT7-2779 with
the senior author. Further syntheses and preparation of this article were supported
by the U.S. Department of Energy, Office of Health and Environmental Research,
under Contract No. DE-AC08-83NV10282 with EG&G Energy Measurements,
Inc. By acceptance of this article, the publisher and/or recipient acknowledges the
U.S. Government’s right to retain a nonexclusive royalty-free license in and to
any copyright covering this paper. Reference to a company or product name does
not imply approval or recommendation of the product by the U.S. Bureau of
Land Management or Department of Energy to the exclusion of others that may
be suitable.

We gratefully thank the following people who contributed to this study: Kristin
Berry of the BLM, Bill Laudenslayer, formerly with the BLM, and John Siperek,
a former BLM employee who assisted with initiation of field work and trained
us in telemetry techniques; Harold J. Egoscue, formerly of the National Zoological
Park, Smithsonian Institution, who also assisted with the first field trip, advised
us on techniques, and identified all fleas; Rory Lamp, Dan Delaney, and Mike
Neitz, our field assistants at various times; Nancy Batson Crews, our helpful,
enthusiastic pilot; Gordon Gould and D. A. Jessup, DVM, of the California
Department of Fish and Game who provided assistance in autopsying a fox; and
Joan T. O’Farrell who provided field assistance in addition to her secretarial skills.

Literature Cited

Anonymous. 1980. Final Environmental Impact Statement and Proposed Plan, California Desert
Conservation Area. U.S. Department of the Interior, Bureau of Land Management, Sacramento,
California. 245 pp + 15 appendices.

ECOLOGY OF THE DESERT KIT FOX 15

Armstrong, W. H., and C.G. Anthony. 1942. An epizootic of canine distemper in a zoological park.
Cornell Vet., 32:286—288.

Beck, D E., and D. M. Allred. 1966. Siphonaptera (fleas) of the Nevada Test Site. Brigham Young
Univ. Sci. Bull., Biol. Ser., 7(2):1—27.

Burt, W. H. 1934. The mammals of southern Nevada. Tran. San Diego Soc. Nat. Hist., 7:375-357.

Egoscue, H. J. 1956. Preliminary studies of the kit fox in Utah. J. Mammal., 37:351-357.

1962. Ecology and life history of the kit fox in Tooele County, Utah. Ecology, 43:481-497.
1975. Population dynamics of the kit fox in western Utah. Bull. So. Calif. Acad. Sci., 74:

122-127.

Goldman, E. A. 1930. The coyote-arch predator. Symposium of predatory animal control. J. Mam-
mal., 11:325-335.

Grinnell, J., J. S. Dixon, and J. M. Lindsdale. 1937. Fur-bearing animals of California, Vol. 2, Univ.
Calif. Press, Berkeley. ix + 777 pp.

Hall, E.R. 1946. Mammals of Nevada. Univ. Calif. Press, Berkeley and Los Angeles. xi + 710 pp.

Huff, G. L., W. J. Bigler, S. J. Proctor, and L. P. Stallings. 1974. Epizootic of canine distemper virus
infection among urban raccoons and gray foxes. J. Wildl. Dis., 10:423-428.

Knapp, D. K. 1978. Effects of agricultural development in Kern County, California, on the San
Joaquin kit fox in 1977. Final Report, Project E-1-1, Job V-1.21, Non-game Wildlife Inves-
tigations, California Department of Fish and Game, Sacramento. 48 pp. + 6 appendices.

Laughrin, L. 1970. San Joaquin kit fox. Its distribution and abundance. Admin. Rept. No. 70-2,
P-R Project W-54-R, California Department of Fish and Game, Sacramento, 20 pp.

Morrell, S. H. 1971. The life history of the San Joaquin kit fox. Unpub. M. S. Thesis, Univ. of

Calif., Santa Barbara. 67 pp.

1972. Life history of the San Joaquin kit fox. Calif. Fish and Game, 58:162-174.

. 1975. San Joaquin kit fox distribution and abundance in 1975. Admin. Rept. No. 75-3, P-R

Project W-54-R-7-1, California Department of Fish and Game, Sacramento. 27 pp.

O'Farrell, T. P., and L. Gilbertson. 1979. Ecological life history of the desert kit fox in the Mojave
Desert of southern California. Final Rept., U.S. Bureau of Land Management, Desert Planning
Staff, Riverside, California. 96 pp.

Schitosky, F. 1975. Primary and secondary hazards of three rodenticides to kit fox. J. Wildl. Mgmt.,
39:416-418.

Scott, T. G., and W. D. Klimstra. 1955. Red foxes and a declining prey population. Monog. Ser.
No. 1, South. Ill. Univ., Carbondale. 123 pp.

Seton, E. T. 1937. Lives of game animals, Vol. 1. Literary Guild, New York. 640 pp.

Sperry, C. C. 1941. Food habits of the coyote. U.S. Dept. of the Interior Wildl. Res. Bull., 4:1-70.

State of California. 1980. At the crossroads. A report on the status of California’s endangered and
rare fish and wildlife. Resources Agency, Fish and Game Commission and Department of Fish
and Game. California Department of Fish and Game, Sacramento. 147 pp.

Swick, C.D. 1973. San Joaquin kit fox—an impact report of secondary hazards of aerial application
of 1080 grain baits for ground squirrel control in San Luis Obispo County. Calif. Dept. of Fish
and Game, Special Wildl. Investigations, Job II-11, Final Report, California Department of
Fish and Game, Sacramento. 14 pp.

Turkowski, F. J. 1974. Fleas of Arizona gray and kit foxes. J. Ariz. Acad. Sci., 9:55.

Young, S. P., and H. H. T. Jackson. 1951. The clever coyote. Stackpole Press, Pennsylvania. 411 pp.

Accepted for publication 15 February 1984.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 16-21
© Southern California Academy of Sciences, 1986

A New Species of Boccardia (Polychaeta: Spionidae)
from the Galapagos Islands and a Redescription
of Boccardia basilaria Hartman from
Southern California

James A. Blake

Battelle New England Marine Research Laboratory, 397 Washington Street,
Duxbury, Massachusetts 02332

A new species of Boccardia is described from shallow subtidal coral heads of
Academy Bay, Santa Cruz Island, Galapagos Islands. The species is a borer and
related to other species from the eastern Pacific. Boccardia basilaria Hartman, a
close relative of the new Galapagos species occurs in subtidal benthic habitats of
the southern California continental shelf. The species is redescribed and several
deficiencies in its original description are corrected.

As revised by Blake and Kudenov (1978), the genus Boccardia Carrazzi is
restricted to spionids having branchiae from setiger 2 and two types of major
spines in the modified setiger 5. Species having only one type of major spine are
now included in the genus Boccardiella Blake and Kudenov.

Most of the seventeen known species of Boccardia have been well described in
a series of papers (Woodwick 1963a, b; Blake 1966, 1979, 1981; Blake and Ku-
denov 1978; Blake and Woodwick 1971; Rainer 1973; Read 1975). Nine species
occur in the eastern Pacific of which eight are endemic.

While examining spionid collections in the U.S. National Museum, an unde-
scribed species of Boccardia was discovered among materials from the Galapagos
Islands collected during the Southeast Pacific Biological and Oceanographic Pro-
gram of 1965-66. The new species, described below, is part of an eastern Pacific
species group having modified spines in the posterior notopodia. While studying
this new form, it was necessary to examine type specimens of the closely related
B. basilaria Hartman from southern California. Several important characteristics
not noted in the original description (Hartman 1961) necessitate a redescription
of that species.

Boccardia galapagense new species
Figure 1

Material examined. —GALAPAGOS ISLANDS, Santa Cruz Island, Academy
Bay, Anton Bruun Sta. 66128, 0°44.50'S; 90°18.35’W, 23 May 1966, rocky point
near Karl Angermeyer’s residence, in coral head on rock near steep dropoff to
muddy-sand bottom with scattered coral heads, 14 m, coll. S. E. Earle and D.
Wallen, SCUBA, holotype (USNM 80487).

Description. — Holotype posteriorly incomplete, large, 22 mm long, 1.2 mm
wide for approximately 120 setigers. Color in alcohol: brown, with dark reticulated
dorsolateral pigment on prostomium.

A NEW SPECIES OF BOCCARDIA 17

aK iy Wy
Wy,

Wg

B
L— 20um alll

30um

L

D

Fig. 1. Boccardia galapagense. A, Anterior end in dorsal view; B, Hooded hook of anterior neu-
ropodium; C, Unhooded hook of posterior neuropodium; D, Capillary notoseta of posterior setiger;
E, Unworn posterior notopodial spine; F, Worn posterior notopodial spine; G, Bristle-topped major
spine from setiger 5; H, Group of two simple, non-bristled and two bristle-topped major spines from
setiger 5.

Prostomium narrow, weakly incised anteriorly, followed by narrow, papillated
medial portion and with folded caruncle continuing to posterior margin of setiger
3; occipital tentacle lacking, but with anterior edge of caruncle partiaily elevated,
overlying anterior portion of prostomium; without eyes; prominent nuchal cilia
present lateral to caruncle (Fig. 1A). Peristomium well developed, slightly inflated;
palps thick, with numerous folds along ciliated groove, extending posteriorly for
about 12 setigers.

Setiger 1 reduced, lacking notopodium and notosetae; neuropodium short, fin-
gerlike (Fig. 1A). Setigers 2-4, 6 with broad, thin, bladelike elliptical postsetal
notopodial lamellae; subsequent notopodia not apparent.

18 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Anterior notosetae and neurosetae of setigers 1—4, 6 short, unilimbate capillaries,
arranged in two rows, with setae of first row shortest, with fine granulations on
shaft and clear sheath; setae of second row longest, lacking granulations. Notosetae
of middle and posterior setigers longer, lacking granulations; posterior notosetae
including 5—6 fimbriated capillaries (Fig. 1D) and 2-3 spines; each spine with
finely bristled, pointed tips when new (Fig. 1E): with blunted, non-bristled tip
when worn (Fig. 1F). Bidentate hooded hooks from neuropodia of setiger 7,
numbering 5-6 at first accompanied by 1-2 very fine capillaries; hooks, increasing
to 7-8 per neuropodium in middle setigers; anterior hooded hooks with large,
expanded hood, having fine striations; with wide angle between main fang and
shaft and about 80° angle between apical tooth and main fang (Fig. 1B); hooks of
posterior setigers numbering 3—4 per neuropodium losing apical teeth and hoods,
becoming straighter and spinelike (Fig. 1C); presence of spines in both noto- and
neuropodia of posterior setigers providing distinct armature.

Modified setiger 5 not heavily muscularized, only slightly larger than preceding
and succeeding setigers (Fig. 1 A); setae including curved row of two types of major
spines and neuropodial fascicle of capillaries; no superior capillary fascicle; major
spines of two types: 1) dorsal row of 7 smooth, non-bristled, spines with slightly
expanded ends; 2) ventral row of 7 bristle topped spines with expanded tips; when
viewed dorsally, these spines with apical concavity open to one side with terminal,
smooth knob; cavity not seen when viewed from ventral side.

Branchiae present on setigers 2—4, 6 and following setigers to posterior end of
fragment. Branchiae of setigers 2—4, 6-7 shortest, increasing to full size by setiger
9 (Fig. 1A); branchiae broad, thin nearly concealing dorsal surface in middle of
body segments; posterior branchiae short, fingerlike. Pygidium unknown.

Remarks.—Boccardia galapagense is closely related to B. pugettensis Blake
from British Columbia and Washington, B. anopthalama Rioja from western
Mexico and B. basilaria Hartman from southern California in having posterior
notopodial spines. The species is closest to B. basilaria in lacking notosetae on
setiger | and in the manner in which bidentate hooded hooks change to unidentate
spinelike setae in posterior setigers. B. galapagense differs from B. basilaria in
having a papillated, narrow, weakly incised prostomium lacking eyes, bristle-
topped major spines of setiger 5 with a distinct apical concavity on the posterior
side of the expanded bristled tip, and posterior unidentate neuropodial spines
lacking hoods. B. basilaria, on the other hand, has a more typical strongly incised
non-papillated prostomium with four eyes, bristles completely cover the expanded
tips of the major spines and the posterior unidentate spines of the neuropodia
bear a distinct hood.

Distribution. —B. galapagense is known only from the Galapagos Islands where
it bores into shallow subtidal coral rock.

Boccardia basilaria Hartman, 1961
Figure 2

Boccardia basilaria Hartman 1961:95—96, pl. 13, figs. 1-7; 1969:89-—90, 5 figs.

Material examined.—CALIFORNIA, 8.7 miles from Santa Barbara Point Light,
Velero Sta. 5161-57, 34°24'35”’N; 119°54'00’W, 2 July 1957, 20.9 m, green mud,
Orange Peel Grab, holotype and 50+ paratypes (AHF Poly 0636-7).

Description. —Holotype complete 18.5 mm long and 1.2 mm wide for 65 se-

A NEW SPECIES OF BOCCARDIA 19

Cc

Ls 20m __J
D F

Fig. 2. Boccardia basilaria. A, Hooded hook from anterior neuropodium; B, Hooded hook from
middle neuropodium; C, Hooded hook from posterior neuropodium; D, Capillary notoseta from
posterior row of anterior setiger; E-F, Spines from posterior notopodia; G, Bristle-topped spine from
setiger 5; H, Simple major spine from setiger 5.

tigerous segments; most paratypes smaller, incomplete. Color in alcohol: light
brown with single medial black pigment spot on 6-10 segments from setiger 5;
middle and posterior segments of some specimens with brown reticulated pigment
on dorsum and similar pigment on prostomium and anterior ventral areas of
some specimens. Body dorsoventrally flattened anteriorly, cylindrical posteriorly.

Prostomium incised anteriorly, forming 2 diverging lobes, continuing poste-
riorly as caruncle to end of setiger 3; no occipital tentacle, but prostomium with
dorsal protuberance at level of setiger 1; with 2 pair of eyes.

Setiger 1 biramous, dorsally elevated; with elliptical neuropodial postsetal la-
mellae and short, fingerlike notopodial lamellae; setigers 2—4, 6 and subsequent
segments with thin, leaflike elliptical noto- and neuropodial postsetal lamellae;
notopodial lamellae of middle segments lower, more elongate; neuropodial la-
mellae less rounded, longer.

Notosetae of setiger 1 simple fascicle of capillaries; setigers 2—4, 6 and following
segments with well-developed fascicles of capillary notosetae arranged in two rows
with setae of first row shorter, thicker, sharply curved with fimbriated edge; setae

20 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

of posterior row longer, straight, without fimbriated edge (Fig. 2D). Posterior
notopodial spines present in posterior setigers; spines simple, numbering 1-2 per
notopodium (Figs. 2E, F), accompanied by 2-4 long, non-limbated capillaries.
Neurosetae of setigers 2—4, and 6 similar in arrangement and form to notosetae.
Most capillaries replaced by hooded hooks from setiger 7; hooks numbering 7-8
in a fascicle, accompanied by 1-3 capillaries in ventralmost position; hooks strong-
ly recurved, with inflated hood and with unusually wide angle between main fang
and shaft, apical tooth reduced and closely adhering to main fang (Fig. 2A); in
middle setigers apical tooth becoming reduced (Fig. 2B) and finally lost in posterior
neuropodia (Fig. 2C); hoods of unidentate hooks in posterior setigers reduced,
closely applied to shaft. Neuropodial hooks and notopodial spines providing
distinct armature to posterior end.

Modified setiger 5 about twice size of preceding and succeeding setigers; setae
including curved row of two types of major spines and neuropodial bundle of
about fifteen thin capillaries with finely granulated shafts and narrow fimbriated
edge; dorsal capillaries lacking. Major spines include dorsal row of 2-3 simple
falcate, non-bristled spines (Fig. 2H) and ventral row of 7-8 bristle topped spines
with bristles completely enclosing tip (Fig. 2G).

Branchiae present on setigers 2—4, 6 and subsequent segments, absent from last
20 or so setigers; all branchiae, short, widely spaced, located closely to notopodium
and nearly fused basally to notopodial postsetae lamellae; anterior branchiae short,
barely extending one-quarter of distance across dorsal midline; branchiae reaching
full size by about setiger 15, extending about half distance to dorsal midline.

Pygidium with two lappets located ventral to dorsally positioned anal opening.

Remarks. —In the original description of B. basilaria, the posterior notopodial
spines were not observed (Hartman 1961), although the unidentate nature of the
posterior neuropodial hooks was described. The close similarity of B. basilaria
and B. galapagense has already been discussed (see above) and relationships with
some other species were considered in an earlier paper (Blake 1979).

Distribution. —Southern California continental shelf.

Acknowledgments

I am grateful to Dr. Kristian Fauchald, formerly of the Allan Hancock Foun-
dation, for the loan of the B. basilaria types. Ms. Susan Williams of the Hancock
Foundation provided station data.

Literature Cited

Blake, J. A. 1966. On Boccardia hamata (Webster), new combination (Polychaeta, Spionidae). Bull.
So. Calif. Acad. Sci., 65:176-185.

1979. Revision of some polydorids (Polychaeta: Spionidae) described and recorded from
British Columbia by Edith and Cyril Berkeley. Proc. Biol. Soc. Wash., 92(3):606-617.
. 1981. Polydora and Boccardia species (Polychaeta: Spionidae) from Western Mexico, chiefly
from calcareous habitats. Proc. Biol. Soc. Wash., 93(4):947-962.
, and J. D. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Australia and
adjacent areas with a revision of the genera. Mem. Nat. Mus. Victoria, 39:171-280.
,and K. H. Woodwick. 1971. Areview of the genus Boccardia Carrazzi (Polychaeta: Spionidae)
with descriptions of two new species. Bull. So. Calif. Acad. Sci., 70:31—42.
Hartman, O. 1961. Polychaetous annelids from California. Allan Hancock Pacific Exped., 25:1-226,

34 pls.

A NEW SPECIES OF BOCCARDIA 21

. 1969. Atlas of the Sedentariate polychaetous annelids from California. Allan Hancock Foun-

dation, Univ. Southern California, Los Angeles, 812 pp.

Rainer, S. 1973. Polydora and related genera (Polychaeta: Spionidae) from Otago waters. Jour. Roy.
Soc. New Zealand, 3:545-564.

Read, G. B. 1975. Systematics and biology of polydorid species (Polychaeta: Spionidae) from Wel-
lington Harbour. Jour. Roy. Soc. New Zealand, 5:395-419.

Woodwick, K. H. 1963a. Comparison of Boccardia columbiana Berkeley and Boccardia proboscidea

Hartman (Annelida, Polychaeta). Bull. So. Calif. Acad. Sci., 62:132-139.

1963b. Taxonomic revision of two polydorid species (Annelida, Polychaeta, Spionidae).
Proc. Biol. Soc. Wash., 76:209-216.

Accepted for publication 16 August, 1984.

Bull. Southern California Acad. Sci.
85(1). 1986, pp. 22-28
© Southern California Academy of Sciences. 1986

Notes on Three European Land Mollusks
Introduced to California

Barry Roth

Santa Barbara Museum of Natural History, 2559 Puesta Del Sol Road,
Santa Barbara, California 93105

Cecilioides acicula (Muller 1774) is newly recorded as an introduction in Cal-
ifornia. Punctum conspectum (Bland 1865) is a synonym of Punctum (Toltecia)
pusillum (Lowe 1831); widely distributed in western North America, it may be
both native and introduced. Previous records in California of Arion circumscriptus
Johnston, 1828, are shown to pertain to Arion (Carinarion) silvaticus Lohmander,
1937: the latest documented finding of the species in the state is 1949.

This paper presents information regarding two species of European terrestrial
gastropods newly recognized as introductions to western North America or to the
state of California and a third which may be both native and introduced. The
records added here supplement those in the summary by Hanna (1966) and
subsequent papers (Roth 1977, 1982a. 1982b; Roth and Chivers 1980) docu-
menting the continuing discovery and spread of synanthropic snails and slugs in
this part of the world.

The following abbreviations are used: CAS—California Academy of Sciences,
San Francisco: RMNH—Rijksmuseum van Natuurlijke Historie, Leiden;
SBMNH~-—Santa Barbara Museum of Natural History.

Cecilioides acicula (Miller 1774)

CALIFORNIA: Santa Clara County, garden in Los Altos, on carrots [Daucus
carota]; J. F. Vedder coll. February 1967 (CAS 036707).

The only previous North American records are in the eastern states— Florida,
Maryland, Pennsylvania, and New Jersey (Pilsbry 1946; Dundee 1974). It has
been intercepted on numerous kinds of plant shipments originating in both New
and Old World ports (Dundee 1974).

The subterranean habit of Cecilioides acicula may account for the scarcity of
records in western North America. The species may in fact have a much wider
distribution in the San Francisco Bay counties, although possibly limited by the
availability of calcium-rich soil. With C. acicula at the above locality was found
Helicodiscus singleyanus (Pilsbry 1890). previously known in the San Francisco
Bay area from roots and bulbs in a garden in Berkeley (Pilsbry 1948).

Punctum (Toltecia) pusillum (Lowe 1831)
Figures 1-2

This species is a familiar member of the fauna of western North America, where
it has gone under the name Punctum (Toltecia) conspectum (Bland 1865). This
synonymy is new herein.

22

INTRODUCED EUROPEAN LAND MOLLUSKS 23

The range attributed to “P. conspectum”’ includes Alaska, British Columbia,
Idaho, Montana, Arizona, and New Mexico, as well as the Pacific Coast states of
Washington, Oregon, and California (Pilsbry 1948; Bequaert and Miller 1973).
It also occurs in Kamchatka (Dall 1905; Likharev 1963). If Punctum jaliscoense
(Pilsbry 1926) is a synonym or a subspecies, as it was treated by Baker (1930)
and Pilsbry (1948), the species ranges to southern Mexico. No fossil occurrences
are known.

I have examined specimens from the following counties in California: Alameda,
Butte, Contra Costa, Humboldt, Los Angeles, Mendocino, Merced, Monterey,
Nevada, Plumas, San Benito, San Bernardino, San Francisco, San Joaquin, San
Luis Obispo, San Mateo, Santa Barbara, Santa Clara, Santa Cruz, Sonoma, Te-
hama, Tulare, and Tuolumne. In addition Pilsbry (1948) cited Calaveras, Mari-
posa, Napa, Orange, San Diego, and Siskiyou counties. Localities are sporadically
distributed through the northern and southern California Coast Ranges, Sierra
Nevada and Cascade Range (principally the foothills), with a few occurrences in
the Transverse and Peninsular Ranges. Many localities are clustered in the San
Francisco Bay area and the Los Angeles Basin. It is not possible to say to what
extent the gaps in range are real or reflect the uneven distribution of collecting
effort.

Specimens in the California Academy of Sciences collection from Alaska, Wash-
ington, Oregon, and Idaho are all very similar in shell characters. Specimens from
Pasadena and adjacent parts of Los Angeles County, formerly regarded as the
subspecies Punctum conspectum pasadenae Pilsbry, 1896, have the major radial
riblets weakly developed (Fig. 1D). Pilsbry (1948) stated that P. c. pasadenae was
usually smaller than the nominate subspecies, but shells in lots I have examined
(Pasadena, R. Arnold coll. [CAS 036854], and H. N. Lowe coll. [CAS 036855];
bed of Los Angeles River near Los Feliz Bridge, Grifhth Park, Los Angeles, M.
E. Caruthers coll. [CAS 036856]) actually average about 10 per cent larger in
diameter at a given whorl than central California specimens.

Specimens from San Francisco, California (the type locality of Punctum con-
spectum) (Fig. 1C), and elsewhere in the west (Figs. 1D-1F) are very simiiar to
authentic specimens of European P. pusillum (Figs. 1A-1B). All have a spirally
lirate protoconch of 1.4 whorls (Figs. 1B, E), 5-10 minor radial riblets in the
intervals between major radials, flattened whorl shoulder, and low spiral cords
corrugating the shell surface (Figs. 2A, B). Typical of Punctum, the radial and
spiral sculpture are essentially independent (Solem 1977).

The recognized range of Punctum pusillum includes the Macaronesian archi-
pelagos and many localities around the Mediterranean, where it has been recorded
most often as Pleuropunctum micropleuros (Paget 1854). Gittenberger et al. (1980)
remarked that its total distribution is poorly known “because the systematic
position of many at least closely related nominal taxa has never been thoroughly
studied: e.g., Helix poupillieri Bourguignat, 1863, from Algeria; Punctum lederi
(O. Boettger 1880), reported from SE. Azerbaydzhan, U.S.S.R. (Lenkoran) to
Israel and Turkey; Patula atoma Gredler, 1892, from China, Hunan province;
Pyramidula javana (Mollendorff 1897) from Java, Bali and Lombok; Punctum
hottentotum (Melvill & Ponsonby 1891) from southern Africa; etc.’’ Based on
a lot from Aomar, Algeria (CAS 036851), I regard Helix poupillieri Bourguignat
as a synonym of P. pusillum. Other material that I find suggestively similar comes

24 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 1. Punctum (Toltecia) pusillum (Lowe). A, B, Spain: prov. Barcelona, Montjuich, El Marrot,
near Miramar, C. Altimira coll. Dec. 1966 (CAS 037053, ex RMNH 54890). C, California: San
Francisco, Buena Vista Park, B. Roth coll. 2 July 1970 (CAS 037054), virtual topotype of Helix
conspecta Bland. D, California: Los Angeles, bed of Los Angeles River near Los Feliz Bridge, Griffith
Park, M. E. Caruthers coll. (CAS 037055). E, F, California: Plumas County, near Blairsden, A. G.

Smith coll. 3 August 1951 (CAS 037056). A, C, D, F, top views, x 25; B, E, sculpture of apical whorls,
x72.

INTRODUCED EUROPEAN LAND MOLLUSKS 25

Pe Lge PF
te Sy yal Sm OFF ya
Oe Re cat

Fig. 2. Punctum (Toltecia) pusillum (Lowe), details of sculpture on shoulder of body whorl, x 345.
A, California: Plumas County, near Blairsden (CAS 037056). B, Spain, prov. Barcelona, Montjuich,
EI Marrot, near Miramar (CAS 037053, ex RMNH 54890).

from Tonglu, Zhejiang province, China (CAS 036857, originally identified as
Punctum orphana [Heude}); and Zhililung, near Pujiang, Zhejiang province, China
(CAS 036858, “P. orphana’’). Dr. E. Gittenberger (written comm. 1983) points
out that Zilchogyra cleliae Weyrauch (1965: pl. 7, fig. 3; not fig. 2 as stated in
text) from Argentina and southeastern Brazil may be yet another synonym. The
type locality of Z. cleliae is a park in Buenos Aires, under the loose bark of
eucalyptus trunks, a characteristic type of habitat for an introduced species.
Throughout much of its North American range, Punctum pusillum is evidently
indigenous: remote localities in Alaska, the mountains of Idaho, Arizona, New
Mexico, and other states. However, it also occurs in some situations that are
strongly suggestive of an introduced species. For example, it was found in asso-
ciation with Vitrea contracta, Helix aspersa, Oxychilus cellarius, and Arion in-
termedius in a landscaped section of San Francisco (Roth 1977); with Helix aperta,
Milax gagates, Deroceras caruanae, and Deroceras reticulatum—all European
species—in Richmond, California (Roth and Chivers 1980). It is a common,
although inconspicuous, inhabitant of parks and gardens in the San Francisco Bay
area. On Santa Cruz Island (southern California) it occurs in a disturbed situation
around the University of California field station (SBMNH 33889) but has not yet
been discovered in native plant communities on this or any other California island.
Other European mollusks, including Milax gagates, Limax maximus, and Limax
valentianus, are similarly restricted to disturbed habitats on Santa Cruz Island.
The far-flung but apparently patchy distribution of Punctum pusillum may be
a consequence of its ‘““weedy”’ tendencies. The species is distinctly anthropophilic,
but the relative deployment of native versus introduced populations and the
source-areas for its urban occurrences will be hard to discover unless some genetic
markers or anatomical differences are found. I believe human introduction from
Europe must continue to be regarded as a possibility, because of the many parallel
instances of Old World mollusks, particularly from the Mediterranean area, be-
coming successfully established in “‘civilized’’ contexts in California. Instances of

26 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

North American species introduced to Europe or the Middle East are far fewer
in number (Mienis 1977; Kerney and Cameron 1979).

Some recent authors (e.g., Falkner 1974; Gittenberger et al. 1980) treat Toltecia
Pilsbry, 1926, as a genus. Toltecia is distinguished from Punctum Morse, 1864,
by the presence of “‘several, usually five or more,” minor radial threads in the
interspaces between major radials on the last whorl (Pilsbry 1948:649). The type-
species of Punctum, P. minutissimum (Lea 1841), has, in general, two or three
such “‘microradials”’ in the interspaces (Solem 1977: figs. 5, 6). This difference is
one of numbers, not of basic structure. In Toltecia the last ¥%3 to % whorl before
maturity enlarges at a greater rate than previous whorls; the inner lip expands
somewhat toward the umbilicus and the plane of the aperture becomes nearly
tangent to the face of the body whorl. In Punctum this terminal change in growth
is absent and the plane of the aperture remains approximately radial (E. Gitten-
berger, written comm. 1983). Species of both Punctum and Toltecia have spiral
liration on the protoconch; that of Toltecia may be slightly stronger. Available
anatomical information does not give grounds for separation; compare Baker’s
(1927: pl. 16, figs. 8, 12) figures of genitalia and pallial organs of Punctum (Toltecia)
jaliscoense (Pilsbry)—the type-species of Tol/tecia—with Solem’s (1983: fig. 24)
of P. minutissimum. Pilsbry (1948) ultimately reduced P. jaliscoense to a sub-
species of Punctum conspectum.

In the absence of a phylogenetic revision of the Punctidae addressing these
relationships on a worldwide basis, to treat Toltecia as a subgenus—as I have
done—actually implies more information than is now at hand.

Arion (Carinarion) silvaticus Lohmander, 1937

CALIFORNIA: San Francisco City and County, Golden Gate Park, near Cal-
ifornia Academy of Sciences buildings, under leaves; A. G. Smith coll. 4 July
1942 (CAS 025655). Same data but coll. 18 January 1947 (CAS 025653). Same
data but coll. 30 April 1949 (CAS 025656).

For many years the distinction between Arion circumscriptus Johnston, 1828,
Arion fasciatus (Nilsson 1823), and Arion silvaticus—first pointed out by Loh-
mander (1937)—was not appreciated by American authors. Most records until
about 1969 simply cite Arion circumscriptus (e.g., Pilsbry 1948; Hanna 1966).
Dissection of the specimens cited above shows them to be A. sil/vaticus. The species
is reported in the New World from northeastern North America (Chichester and
Getz 1969; Dundee 1974) and from British Columbia (Rollo and Wellington
1975); this is the first record from California.

These specimens were mentioned by Hanna (1966:29) as A. circumscriptus.
Although 4A. silvaticus persisted in this locality from at least 1942 to 1949, it had
disappeared by the time of Hanna’s (1966) report. I have not been able to find it
subsequently. Lange (1944) collected “‘A. circumscriptus’’ in Golden Gate Park
in 1940 and quoted A. G. Smith (in correspondence) that the species was wide-
spread in the San Francisco Bay region. I have not managed to track down any
of Lange’s specimens, and the collection of the California Academy of Sciences,
which contains most of the slug material handled by A. G. Smith, includes no
Bay Area lots beside those cited above.

The several additional reports of Arion circumscriptus from California (Gregg
1943; Pilsbry 1948; Ingram 1949; Ingram and Lotz 1950) seem to be restatements

INTRODUCED EUROPEAN LAND MOLLUSKS 27

of the same information, referencing either the 1944 Lange publication or personal
communication from Smith. The statement by Hill (1951) that A. circumscriptus
occurs in greenhouses in the San Francisco Bay area is probably also derivative.
In summary, although the lots listed here document the former presence of Arion
silvaticus in this part of California, evidence for its wider occurrence is equivocal
or lacking and its continued presence remains problematical.

Acknowledgments

For circumstances permitting field work on Santa Cruz Island, California, I am
indebted to Dr. F. G. Hochberg, Curator, Department of Invertebrate Zoology,
SBMNH, Dr. Carey Stanton, and the Santa Cruz Island Company. Field station
facilities were made available through the Santa Cruz Island Reserve administered
by the University of California. I thank Dr. Edmund Gittenberger, Curator, De-
partment of Mollusca, RMNH, for lending authentic specimens of Punctum (Tol-
tecia) pusillum to compare with Californian material and for commenting on an
earlier draft of the manuscript. The scanning electron micrographs were taken at
the Electron Microscope Facility of San Francisco State University, kindly made
available by Dr. Stanley C. Williams and Mr. Kenneth Letch.

Literature Cited

Baker, H. B. 1927. Minute Mexican land snails. Proc. Acad. Nat. Sci. Philadelphia, 79:223-246.
1930. Mexican mollusks collected for Dr. Bryant Walker in 1926. Part II. Auriculidae,

Orthurethra, Heterurethra, and Aulacopoda. Occas. Pap. Mus. Zool. Univ. Michigan, 220:1—55.

Bequaert, J. C., and W. B. Miller. 1973. The mollusks of the arid southwest. Univ. Arizona Press,
xvi + 271 pp.

Chichester, L. F., and L. L. Getz. 1969. The zoogeography and ecology of arionid and limacid slugs
introduced into northeastern North America. Malacologia, 7:313-346.

Dall, W.H. 1905. Land and fresh water mollusks of Alaska and adjoining regions. Harriman Alaska
Ser., 13:1-171.

Dundee, D. S. 1974. Catalog of introduced molluscs of eastern North America (north of Mexico).
Sterkiana, 55:1-37.

Falkner, G. 1974. Uber Acanthinulinae aus dem Obermiozin Siiddeutschlands (Gastropoda: Pupil-
lacea). Arch. f. Molluskenk., 104:229-245.

Gittenberger, E., H. P. M. G. Menkhorst, and J. G. M. Raven. 1980. New data on four European
terrestrial gastropods. Basteria, 44:11-16.

Gregg, W. O. 1943. Other species of land slugs found in the Pacific Coast states. Min. Conchol. Club
South. California, 27:3-9.

Hanna, G D. 1966. Introduced mollusks of western North America. California Acad. Sci. Occas.
Pap., 48:1-108.

Hill, H. R. 1951. Exotic Mollusca in California. Min. Conchol. Club South. California, 107:1-4.

Ingram, W. M. 1949. A check list of the Limacidae, endodontidae, Arionidae, Succineidae, Pu-
pillidae, Valloniidae, Carychiidae, and Truncatellidae of California. Bull. South. California Acad.

Sci., 48:19-34.

, and C. Lotz. 1950. Land mollusks of the San Francisco Bay counties. Jour. Entomol. Zool.

(Pomona Coll.), 42:20-39.

Kerney, M. P., and R. A. D. Cameron. 1979. A field guide to the land snails of Britain and north-
west Europe. William Collins Sons and Co., 288 pp.

Lange, W. H. 1944. Land slugs in California. Bull. South. California Acad. Sci., 43:33-40.

Likharev, I. M. 1963. [On the fauna of the terrestrial mollusks in the Kamchatkan region.] Pp. 65—
81 in [The fauna of the Kamchatkan region]. (L. A. Portenko, ed.), Akad. Nauk SSSR [In
Russian].

Lohmander, H. 1937. Uber die nordischen Formen von Arion circumscriptus Johnst. Acta Soc. Pro.
Fauna Flora Fenn., 60:90-112.

28 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Mienis, H. K. 1977. North American land snails in Israel. Nautilus, 91:30-31.
Pilsbry,H. A. 1946. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Philadelphia,
Monogr. 3, 2(1):1-—520.

1948. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Philadelphia,
Monogr. 3, 2(2):521-1113.

Rollo, C. D., and W. G. Wellington. 1975. Terrestrial slugs in the vicinity of Vancouver, British
Columbia. Nautilus, 89:107—-115.

Roth, B. 1977. Vitrea contracta (Westerlund) (Mollusca: Pulmonata) in the San Francisco Bay area,
California. Veliger, 19:429-—430.

—. 1982a. European land mollusks in the San Francisco Bay area, California: Carychium

minimum Miiller and the Arion hortensis complex. Veliger, 24:342-344.

1982b. Discus rotundatus (Miller) (Gastropoda: Pulmonata) in California. Malacol. Rev.,
15:139-140.

—., and D. D. Chivers. 1980. Helix aperta introduced in Richmond, California (Mollusca:
Pulmonata). Veliger, 22:385-—387.

Solem, A. 1977. Shell microsculpture in Striatura, Punctum, Radiodiscus, and Planogyra (Pulmo-

nata). Nautilus, 91:149-155.

1983. Endodontoid land snails from Pacific Islands (Mollusca: Pulmonata: Sigmurethra).
Part II. Families Punctidae and Charopidae, Zoogeography. Field Museum of Natural History,
1x + 336 pp.

Weyrauch, W. K. 1965. Neue und verkannte Endodontiden aus Stidamerika. Arch. f. Molluskenk.,
94:121-134.

Accepted for publication 25 July 1984.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 29-34
© Southern California Academy of Sciences, 1986

The Taxonomic Status of Two North American Lithurge
(Hymenoptera: Megachilidae)

Roy R. Snelling

Natural History Museum of Los Angeles County,
900 Exposition Boulevard, Los Angeles, California 90007

Abstract. —Subsequent to my 1983 review of the North American species of
Lithurge, the type specimen of L. planifrons (Friese) has been examined: it is a
senior synonym of L. socorroensis (Mitchell) (NEW SYNONYMY). The species
that I incorrectly believed to be L. planifrons is here described as L. bitorulosa,
no prior name being available. A corrected key to the North American species is
included.

In 1983 I reviewed the North American species of the bee genus Lithurge.
Including one introduced Palearctic species, a total of eight species were recog-
nized. A few Mexican specimens from Jalisco were tentatively identified as L.
planifrons (Friese), a species otherwise known only from the inadequately de-
scribed male type. That type, in the collections of the Zoologisches Museum,
Humboldt-Universitat, Berlin, has been examined at my request by T. L. Gris-
wold. With the aid of Griswold’s notes I have been able to determine that the
specimens I had referred to this name were incorrectly identified, and that Friese’s
name is a senior synonym of L. socorroensis (Mitchell). Since no prior name is
available for the species that I had called L. planifrons it is here named and
described as L. bitorulosa.

A new key is given to reflect these changes. And, the opportunity is taken to
improve the separation of females of L. echinocacti and L. planifrons (L. socor-
roensis in my previous key).

Lithurge (Lithurgopsis) planifrons (Friese)

Lithurgus planifrons Friese, 1908:62. 6.
Lithurgus socorroensis Mitchell, 1938:152-154. 2. NEW SYNONYMY.
Lithurge (Lithurgopsis) socorroensis: Snelling, 1983:3, 10. 8.

The type of L. planifrons is in the Berlin Museum and was examined in 1984
by Griswold. His notes on the type make it clear that the male type is not different
from males of L. socorroensis. It is also clear that the specimens from Jalisco that
1 (1983) recorded as L. planifrons are not that species, but represent one previously
undescribed.

The type locality of L. planifrons is given as ““Jacubaya, Mexico,” a misprint
for Tacabaya, a suburb of Mexico City. As the name implies, L. socorroensis was
based on material from Socorro Island in the Revillagigedo Archipelago. The
range of L. planifrons extends from southern Arizona to Costa Rica.

29

30 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ZN
Ves)
lo. Q 3

2

Figs. 1-4. Lithurge spp., females: 1, L. bitorulosa, frontal view of lower face; 2, L. bitorulosa,
dorsal view of supraclypeal prominence; 3, L. planifrons, same; 4, L. echinocacti, same. Stippling
indicates extent of punctate areas in Figures 2—4. All Figures drawn to same scale.

Lithurge (Lithurgopsis) bitorulosa, new species
Figs. 1-2

Lithurge (Lithurgopsis) planifrons: Snelling, 1983:2, 10. 6. [Misidentification].

Diagnosis. — Female: First flagellar segment shorter than second; supraclypeal
area with large conical protuberance on each side; malar area linear, without
posterior pit; upper facet of supraclypeal process conspicuously contiguously to
subcontiguously punctate. Male: Tarsal arolia absent; labrum with transverse basal
ridge, but without erect spine-like tubercle; supraclypeal area flat, shiny to slightly
shiny between contiguous to subcontiguous punctures.

Description.—Female, Measurements (holotype in parentheses): head width
3.54-4.51 (4.51); head length 2.36—-2.87 (2.87); wing length 7.5—10.0 (10.0); total
length 11.5-15.5 (15.5) mm.

Head: 1.3—1.4 times broader than long; eyes strongly convergent below, upper
interocular distance 1.5—1.6 times lower interocular distance. Malar space essen-
tially absent, posterior pit absent. Transverse basal ridge of labrum high in center,
with deep median excision. Clypeus with narrow, curved preapical ridge, distad
of which clypeus is depressed, shiny; clypeal disc shiny, subpolished between
coarse very variably spaced punctures, but with interspaces up to three times

NORTH AMERICAN LITHURGE 31

puncture diameter in middle. Supraclypeal area moderately protuberant, with
prominent blunt, conical protuberance on each side; lower facet polished and
virtually impunctate, upper facet (including area between bases of protuberances)
dull to moderately shiny between contiguous to subcontiguous, moderate to coarse
punctures that are continuous across midline. Ocellocular distance about 1.3 times
interocellar distance. First flagellar segment shorter than pedicel, about as long
as second segment; median flagellar segments about one-third broader than long.

Mesosoma: Dorsal portion of mesoscutum coarsely, transversely scabrous an-
teriorly, becoming more finely, less distinctly transversely scabrous distad. Disc
of scutellum finely scabrous.

Pilosity: Hairs mostly whitish; ferruginous hairs present on lower margin of
mandible, middle one-third of apical margin of clypeus, and inner side of pro-
basitarsi; meso- and metabasitarsi with brownish hairs on inner side. Sternal scopa
somewhat brownish yellow. Marginal pubescent fasciae of metasomal terga 2 and
3 broadly interrupted, those of 4 and 5 complete; tergum 6 wholly covered with
dark brownish hairs, the apices of which may be lighter reddish brown; tergal
disc with sparse, suberect to erect, simple, brown hairs.

Color: Uniformly blackish brown, but legs (especially femora) lighter, flagellum
ferruginous. Wings transparent, weakly brownish.

Male, Measurements: head width 3.64—3.79; head length 2.97—3.08; wing length
7.5-9.0; total length 11.0-13.0 mm.

Head: 1.2—1.3 times broader than long; eyes strongly convergent below, upper
interocular distance 1.5-1.6 times lower interocular distance. Labrum with low,
rounded basal ridge impressed in middle, but without erect spine. Supraclypeal
area mostly flat, but lower portion sloping toward base of clypeus, 1.4—1.5 wider
than long; clypeus, supraclypeal area slightly shiny between contiguous to sub-
contiguous, moderate to coarse punctures, without impunctate median areas.
Ocellocular distance about 1.3 times interocellar distance; ocelloccipital distance
about 1.3 times interocellar distance. First flagellar segment conspicuously broader
than long, shorter than pedicel and about one-third as long as second segment.

Mesosoma: Mesoscutum with moderate, contiguous punctures, sometimes sub-
rugosopunctate. Scutellum with moderate, contiguous punctures.

Pilosity: Similar to that of female, but without ferruginous hairs on mandible
and clypeal margins, with yellowish ferruginous hairs on inner side of basitarsi;
some brown hairs across occiput and on mesoscutum; metasomal terga 4-6 with
continuous distal fasciae, brown hairs of tergal discs longer, last tergum with hairs
sparse, some pale hairs at sides.

Color: As described for female, but legs definitely brown.

Type material.—Holotype female, allotype, five female paratypes: 46 mi W
Tehuantepec, 2125 ft elev., Oaxaca, MEXICO, 26 June 1961 (University of Kan-
sas Mexican Expedition), on cactus, Opuntia. Types in collection of University
of Kansas, except one paratype in Natural History Museum of Los Angeles County.

Additional material (Not paratypes).—2 6, Estacion Biologia UNAM “Cha-
mela,” San Patricio, Jalisco, MEXICO, 1981 (S. H. Bullock; #76); 1 6, same data
except 21 June 1983 (#1533), in LACM and Estacion Biologia UNAM “Chamela.”’

Etymology.—Combines the Latin prefix ‘“bi-”’ (two) with the diminutive of
torus (protuberance or bulge).

Discussion. —The female of L. bitorulosa will run to the last couplet in my 1983

32 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

key, separating L. echinocacti (Cockerell) and L. planifrons (as L. socorroensis).
In all three of these species the supraclypeal area has a pair of sublateral conoid
processes. The female of L. echinocacti has the supraclypeal area without defined
upper and lower facets and with most of its area smooth and shiny between
scattered fine punctures; only on the area adjacent to the antennal sockets do the
punctures become crowded; the supraclypeal tubercles, in dorsal view, are narrow
and acute and the entire supraclypeal area is less strongly protuberant (Fig. 4).
The supraclypeal structure of L. planifrons more closely resembles that of L.
bitorulosa, but the shiny, impunctate lower facet extends up and includes all of
the area between the tubercles (Fig. 3), while in L. bitorulosa the lower facet is
confined to the lower one-third of the supraclypeal area, with most of the area
between the tubercles subcontiguously punctate (Fig. 2). In both L. echinocacti
and L. planifrons, metasomal terga 2 and 3 possess apical hair bands that are
complete across the middle of the segments.

The male, misidentified by me as L. planifrons in 1983, will run to that name
in my key. Because the labrum lacks an erect spine-like process, the male is most
like those of L. echinocacti and L. apicalis. It differs from both in having the
supraclypeal area flat and coarsely and closely punctate along the midline.

The following is a revised version of my 1983 key, reflecting the changes made
herein.

Key to North American Lithurge

lay Female} metasoma with sixiexposedstencal oa. 4) eee 2

b. Male, metasoma with seven exposed terga ........................ 9
2a. First flagellar segment shorter than second; facial prominence, when

present, not as below (subgenus Lithurgopsis) ..................... 3

b. First flagellar segment longer than second; facial prominence high, tri-
angular in lateral view, its lower portion sloping toward base of clypeus

Gubgenusy/oihunce) eee eee chrysurus (Fonscolombe)
3a. Supraclypeal protuberance nearly as broad as face, extending laterad of
subantennal sutures; malar area distinct and with deep posterior pit .. 4

b. Supraclypeal protuberance, when present, much narrower than width
of face, not extending laterad of subantennal sutures; malar area linear,

mandible nearly contiguous with eye, posterior pit usually absent ... 5

4a. Labral tubercle divided in middle only; clypeus with preapical ridge
whichhissinterruptedimni middle sere ee eee apicalis (Cresson)

b. Labral tubercle with median and sublateral impressions; clypeus with-
out preapicalsridgen. wie AAR he et ee eee littoralis (Cockerell)

5a. Supraclypeal area distinctly elevated on each side of midline or with a
single; transverse, bowedielevation’ 445-255-200 e eee 6

b. Supraclypeal area flat, with no protuberance, shiny and very sparsely
punctate: «et. FRel eae AK eS Jo eee Dee hese listrota Snelling

6a. Supraclypeal area with prominent paired, cone-like elevations; malar
area: without: posterior: pit. ws So ee eee 7

b. Supraclypeal elevation a high, transverse, bowed ridge; malar area with
broadyshallowaposteriompitye) oe eee eee gibbosa (F. Smith)

7a. Supraclypeal area very shiny and largely impunctate over one-half or

NORTH AMERICAN LITHURGE 33

8a.

13a.

14a.

15a.

more of its area (in particular, area between bases of lateral tubercles
smooth and shiny); metasomal terga 2 and 3 with complete fasciae ... 8
Supraclypeal area mostly contiguously punctate, only narrow lower
facet impunctate and shiny; metasomal terga 2 and 3 with apical fasciae
broadly interrupted across middle ............... bitorulosa new species
In dorsal view, supraclypeal area only moderately elevated, and lateral
tubercles narrow and acute (Fig. 4); ocelloccipital distance less than
IME ROCE Mare GIS tances hers aa as ee echinocacti (Cockerell)
In dorsal view, supraclypeal area strongly elevated and lateral tubercles
bluntly conoid (Fig. 3); ocelloccipital distance greater than interocellar

GUIS PAIN COMMER REAPS ste I ii gs a A Ee aa planifrons (Friese)
. Tarsal arolia present; first flagellar segment shorter than second (sub-
SEIMUIS MIELE TE COD STS) Harwich ry REE cel cece. ec 2 Nin aa AB see de ce e ae 10
Tarsal arolia absent; first flagellar segment distinctly longer than second
(SUDEEIMUSBILITNULCE) Mn me nies e cs oe chrysurus (Fonscolombe)
. Labrum with a single median tubercle, or none ................... 11
Labrum with a pair of slender, erect tubercles ....... littoralis (Cockerell)
Labrum with a low, transverse basal ridge or swelling .............. 12
Labrum with a slender, erect, spine-like process ................... 14

. Supraclypeal area gently convex, its upper portion sloping toward an-

tennal sockets, and with a definite median area that is less closely
punctate than area near subantennal suture or entire supraclypeal area
SpanselyeandsimnesularlyapumMctatemunan ea irae ce ren) areas 13
Supraclypeal area uniformly flat between clypeal base and antennal
sockets, contiguously punctate, punctures coarse and subcontiguous
AlOn cen AiwMSs Me et et eee MSS Se at Lh. bitorulosa new species
First flagellar segment, on shortest side, distinctly broader than long,
shorter than pedicel; supraclypeal area about 1.4 times wider than long;
lessausualliy sneddishir, yc See als Sed OAR echinocacti (Cockerell)
First flagellar segment, on shortest side, about as long as wide, about
as long as pedicel; supraclypeal area about 1.8 times wider than long;

LESH LOW IW Re Ie MeO tC Ee Oe Uae tne er apicalis (Cresson)
Labrum with low, often obsolescent, ridge extending obliquely from
basaleconnenmtorbasc Olemedianitubercle’ = 5455545. see ane ee WS)

Labrum deeply concave on either side of tubercle and with short, curved
ridge from base of tubercle to midlength of lateral margin ..........

9b. otel adel hte, Bic Jk caes Gore ck Ae Re ML Og planifrons (Friese)
Supraclypeal area slightly protuberant, densely punctate; ocelloccipital

distance greater than interocellar distance ........... gibbosa (F. Smith)

Supraclypeal area flat, sparsely punctate, at least in middle; ocelloccip-

ital distance less than interocellar distance .............. listrota Snelling
Acknowledgments

I wish to thank S. H. Bullock and C. D. Michener for making available the
Mexican specimens that are the basis for this paper. Particular gratitude is ex-
pressed to T. L. Griswold whose examination of the type of L. planifrons at the
Berlin Museum enabled me to correct the errors in my previous paper. I am also

34 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

indebted to R. W. Brooks who had also examined the L. planifrons type and
advised me that my earlier interpretation was incorrect.

Literature Cited

Friese, H. 1908. Die Apidae (Blumenwespen) von Argentina nach den Reisenergebissen der Herren
A. C. Jensen-Haarup and P. Jorgensen in den Jahren 1904-1907. Deutsche Entomologische
Zeitschrift, 1908:1—94.

Mitchell, T. B. 1938. The bee genus Lithurgus (Hymenoptera: Megachilidae). Psyche, 45:146-155.

Snelling, R.R. 1983. The North American species of the bee genus Lithurge (Hymenoptera: Megachil-
idae). Contributions in Science, Natural History Museum of Los Angeles County, no. 343, 11
pp.

Accepted for publication 23 October 1984.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 35-45
© Southern California Academy of Sciences, 1986

Trace Element Composition of Obsidian Butte,
Imperial County, California

Richard E. Hughes

Anthropological Studies Center, Sonoma State University,
Rohnert Park, California 94928

Abstract. —Non-destructive energy dispersive x-ray fluorescence analyses were
performed on rhyolitic volcanic glass (obsidian) from Obsidian Butte, located at
the south end of the Salton Sea in Imperial County, California. Sixteen trace and
rare earth element concentrations were measured in parts per million (ppm), and
these values compared favorably with analyses employing destructive techniques.
The results of the present study demonstrate that archaeological artifacts can be
matched reliably to the Obsidian Butte source on the basis of non-destructive
quantitative analysis, facilitating the study of prehistoric Southern California ob-
sidian trade and interlaboratory comparison of analytical results.

Obsidian has long been of interest to archaeologists because in regions where
it occurs it was the raw material preferred for the manufacture of prehistoric
chipped stone tools. Recently, techniques from the physical sciences have provided
new ways of deriving information from obsidian artifacts. Using x-ray fluorescence
or neutron activation techniques, it is possible to determine the trace and rare
earth element composition of volcanic glass (obsidian), and to distinguish between
parent geological sources on the basis of contrasts in these elemental constituents.
Once the trace element configurations (or “profiles’’) for obsidian sources in a
region are established, it is possible to analyze artifacts from archaeological sites
and match them with known obsidian sources on the basis of trace and rare earth
element similarities. This procedure, popularly known as “fingerprinting,” has
been profitably applied to studies of prehistoric trade in western North America
(e.g., Jack and Carmichael 1969; Jackson 1974; Jack 1976; Hughes 1983a, b,
1984; Ericson 1981; Nelson 1984).

However, until quite recently, comparatively little research on obsidian trade
has been conducted in southern California in part because of the absence of a
published quantitative obsidian source data base. Trace and rare earth concen-
tration values have been published for some elements (see Ericson 1981:10; Rob-
inson et al. 1976:table 2), but these have been derived from destructive analyses
(i.e., those necessitating crushing and powdering some portion of the sample),
which have obvious drawbacks for archaeological studies because of the desir-
ability of analyzing artifacts without sacrificing some portion of them for study.
The purposes of this study are to present a series of quantitative trace and rare
earth element measurements for Obsidian Butte glass using non-destructive x-ray
fluorescence, to compare these values with those generated from previous analyses
conducted using destructive methods, and to illustrate the benefits of non-

35

36 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ACOSO HOT anne ens
SPRINGS Gant

CALIFORNIA

Barstow
©

« \Salton Sea
eS © SDi-682 <2)
SP SDi-5130 <A OBSIDIAN
PAGHEIGH= a. OsDi-813 BUTTE
OCEAN .2=) ©SDi-6071

SanDiegoR W049. ———

ee eGo

Fig. 1. Map of southern California showing the locations of obsidian sources utilized during
prehistoric times (open triangles) and archaeological sites (filled dots) discussed in the text.

destructive quantitative analysis for the study of prehistoric obsidian trade in
southern California.

The Setting

The presence of rhyolitic obsidian at Obsidian Butte has been known to geol-
ogists for some time (e.g., Rogers 1934; Kelley and Soske 1936). However, Ob-
sidian Butte (Fig. 1) was first brought to the attention of the archaeological com-
munity by Treganza (1942:155, fn. 13) and Heizer and Treganza (1944:305), who

TRACE ELEMENTS OF OBSIDIAN BUTTE 37

noted abundant evidence for its use during prehistoric times as a quarry source
for obsidian tools. More than two decades later, Ericson et al. (1976:222) provided
a more detailed description of the immediate geological context. Obsidian Butte
is actually only one of four volcanoes (Obsidian Butte, Red Island, Rock Hill,
and Mullet Island) in the vicinity of the Salton Buttes (see Robinson et al. 1976:
fig. 2) where Quaternary volcanic activity produced rhyolitic obsidian and pumice.
However, petrochemical analyses indicates no systematic elemental variation
between these rhyolitic domes (Robinson, et al. 1976:354). K-Ar ages of ca. 16,000
B.P. (Muffler and White 1969:162) and 33,000 + 35,000 B.P. (2 c) (Friedman
and Obradovich 1981:40—41) have been reported for Obsidian Butte obsidian.

Methods

To determine whether non-destructive analysis could yield quantitative ele-
mental measurements comparable to those previously published using more time
consuming, destructive techniques, analyses were conducted on a sample of 10
unmodified obsidian flakes collected from the southeast side of Obsidian Butte
in T11S, R13E, NE % of the NW % of the SE % of Section 32, as depicted on the
USGS Obsidian Butte, Calif., 7.5’ series quadrangle (1956). No special sample
preparation was made, other than rinsing each specimen in distilled water to
remove surface contaminants that might effect the analysis.

Laboratory analyses were conducted by the author at the Department of Geology
and Geophysics, University of California, Berkeley, on a Spectrace 440 (United
Scientific Corporation) energy dispersive x-ray fluorescence machine equipped
with a 572 power supply (50 kV, 1 mA), 534-1 pulsed tube control, 588 bias/
protection module, 514 pulse processor (amplifier), Tracor Northern 1221 100
mHz analog to digital converter, a Tracor Northern 2000 computer based analyzer
with a LSI-11 microcomputer with 24k word capacity, and a Si(L1i) solid state
detector with 142 eV resolution (FWHM) at 5.9 keV in a 30 mm/? area. Trace
element analyses in the 5-25 keV region of the energy spectrum were conducted
using a Rh x-ray tube operated at 30.0 kV, .40 mA pulsed with a .04 mm Rh
primary beam filter, while analyses in the 20-60 keV region were conducted using
an Am241 100 mCi radioisotope (see Hughes 1983a:fig. 2). All analyses were
conducted at 200 seconds livetime. NI (Ka), Cu (Ka), Zn (Ka), Ga (Ka), Pb (L8),
Th (La), Rb (Ka), Sr (Ka), Y (Ka), Zr (Ka), and Nb (Ka) analytical lines were
excited by the Rh x-ray tube, while Ba (Ka), La (Ka), Ce (Ka), Pr (Ka), and Nd
(Ka) lines were analyzed using the Am241 source. Following excitation, overlap-
ping Ka, K@, and L-lines were stripped, yielding the “‘fingerprints” that appear
in Figure 2 (cf. Hampel 1984). Background-subtracted net peak intensities were
compared and ratioed to the Rh Ka peak continuum (Franzini et al. 1976), then
converted to parts per million using a least squares polynomial fit routine (Scham-
ber 1977; Bice 1980:421—422; Hughes 1983a:25-26).

Results

Most of the quantitative trace and rare earth elemental measurements generated
from the present non-destructive study (Table 2) are in agreement with those
derived from previous destructive analyses (Table 4), and with recommended
values for U.S. Geological Survey international rock standards (Table 1). Pr, Cu,

38 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Zr Rh
5000 (Ko) Ke)
ZEA
4000 (KB) Rh
(KS)
3000 \/ Nb
= (Ko) (Ke)
=
3 Rb
2000 (Kz) LT=200 Seconds
or
1000 (Ka)
0
11.900 13.900 15.900 17.900 19.900 21.900
ENERGY (KeV)
a
Rh
5000 Zr i)
(K2)
4000
YW 3000 Zr
Z (Ks) Rh
5 Y KB
© 2000 (Ke) Nb
Rb (Ka)
(Ka) LT=200 Seconds
1000 Si
(K2)
)
11.900 13.900 15.900 17.900 19.900 21.900

ENERGY (KeV)

b

Fig. 2. Energy dispersive x-ray fluorescence “fingerprint” of Obsidian Butte volcanic glass. a)
‘fingerprint’ of pressed powder sample OS-21 (see Table 4); b) “‘fingerprint’’ of unmodified obsidian
flake sample OB-8 (see Table 2).

and Ni are not measured well by non-destructive x-ray fluorescence, because
concentrations for these elements in obsidians often fall below the minimum
limits of detection (Vane 1977) at 200 seconds livetime. Computation of coeffi-
cients of variation for sample means and standard deviations (Table 3) shows
that these measurements vary widely across samples. Th, Nb, and Sr measure-

39

TRACE ELEMENTS OF OBSIDIAN BUTTE

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42 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Table 4. Trace and rare earth element concentrations for obsidian (OS-21) and whole rock rhyolites
from Obsidian Butte. Obsidian Butte and Red Island data from Robinson et al. (1976: table 2); n.m. =
not measured: = R. N. Jack, analyst. All values in parts per million.

Obsidian Butte Red Island
Element OS-212 170-4A W-772 170-26A 64CMp-107 + 170-21p

Th 26 n.m. n.m. n.m. n.m. n.m.
Pb 16 — = 15 — 10
Nd 62 30 _ 70 — 70
Pr 15 n.m. n.m. n.m. n.m. n.m.
Ce 132 150 — 100 — 150
La 72 70 70 70 30 100
Ba 467 300 700 500 700 500
Nb 31 30 20 20 15 30
Zr 324 300 300 300 300 300
Ye 57 100 100 100 100 150
Sr 30 727] 50 33 50 25

Rb 144 146 n.m. 137 n.m. 142
Ga 21 30 30 30 50 20
Zn 68 n.m. n.m. n.m. n.m. n.m.
Cu 6 3 7 7 10 7
Ni 20 n.m. n.m. n.m. — —

Co 36 — — ES — —

Mn 431 n.m. n.m. n.m. n.m. n.m.
Ti 1074 n.m. n.m. n.m. n.m. n.m.

ments also are more variable than Pb, Nd, Ce. La, Ba, Zr, Y, Rb, Ga, and Zn,
suggesting that they may not be as useful as those remaining elements in distin-
guishing between obsidian sources (see Hughes 1982:176). However, high “ap-
parent” measurement error (relatively high CV% values) will be registered when
the absolute concentration for an element approaches its minimum limits of
detection, and when “true” intersource elemental variability exists. In short, it is
important to recognize that both instrumental measurement limitations and in-
herent geochemical variability may be represented in CV% values.

Obsidian Butte trace element concentrations for Ce, Ba, Zr, Y, Sr, and Rb
contrast with those for other southern California obsidians (Coso Hot Springs
[Jack 1976:table 11.5; Bacon et al. 1981:table 3] Rustler Canyon, San Bernardino
County) and Arroyo Matomi, Baja California. These differences between sources
can be illustrated using a bivariate scatter diagram (Fig. 3).

In order to test whether the results of the Obsidian Butte analyses could prof-
itably be applied to archaeological research, a sample of 52 obsidian artifacts from
five archaeological sites in San Diego County was analyzed and the results were
compared to the trace element profiles generated for the known sources in the
southern California region (see Fig. 3). As Figure 3 shows, these data fall within
the range of trace element concentrations for source specimens from southern
California, showing that 39 artifacts match the profile of Obsidian Butte volcanic
glass, while the remaining 13 derive from the Coso Hot Springs source. Although
these artifacts make up only a part ofa larger study of San Diego County obsidian
trade (see Hughes and True 1985), they illustrate clearly the potential for em-
ploying non-destructive quantitative analyses in archaeological research.

TRACE ELEMENTS OF OBSIDIAN BUTTE 43

150
fe) oe
° ©
125
e oe Fr8 OBSIDIAN
© ee oo ® BUTTE
out bf
100 @
ee te A
4 A
mM A
(pp ) 5 A
COSO HOT
SPRINGS 2% F
ARROYO
A MATOMI
RUSTLER
CANYON
A
0 ie ap ote oa nn ars
100 200 300 400

Zr (ppm)

Fig. 3. Scatter diagram of Y vs. Zr concentrations for obsidian source specimens (open triangles)
from southern California and Baja California. Circles represent values for artifacts from archaeological
sites SDi-682 (@); SDi-813 (©); SDi-5130 (®); SDi-6071 (©) and W-1049 (@). Obsidian Butte source
standards plotted from data in Table 2; Rustler Canyon and Arroyo Matomi (Baja California) values
determined by R. N. Jack using wavelength dispersive x-ray fluorescence.

More generally, the agreement between the present analysis and previous de-
structive work shows that non-destructive analyses can yield concentration esti-
mates (in parts per million) suitable for use in interlaboratory comparisons. The
advantages of reporting the results of archaeometric research in standard mea-
surement units (e.g. weight percent or parts per million) have been recognized for
some time (see Bieber et al. 1976); the results of the present study indicate that
it is now possible to begin building a quantitative geochemical data base for
obsidian source specimens and archaeological artifacts from southern California
that can readily be used by researchers at other laboratories.

Acknowledgments

I thank J. H. Hampel (Department of Geology and Geophysics, University of
California, Berkeley) and J. L. Blomberg (Palo Alto Research Center, Xerox Cor-
poration) for reviewing and commenting on the manuscript; P. J. Wilke (De-
partment of Anthropology, University of California, Riverside) for providing
Obsidian Butte source samples; D. L. True (Department of Anthropology, Uni-
versity of California, Davis), M. Sampson (California Department of Parks and
Recreation), J. Thesken and D. Quillen (WESTEC Services, Inc.) for providing
artifacts for analysis; T. Ruhstaller (Haggin Museum, Stockton) for graphics; and
R. N. Jack (Silverton, Oregon) for generously allowing me to report the results of
some of his previous obsidian analyses.

44 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

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, 1. A. Hagan, and C. W. Chesterman. 1976. Prehistoric obsidian in California II: geologic

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, and D. L. True. 1985. Perspectives on the distribution of obsidians in San Diego County,

California. North Amer. Archaeol., 6:325-339.

Jack, R.N. 1976. Prehistoric obsidian in California I: geochemical aspects. Pp. 183—217 in Advances

in obsidian glass studies: archaeological and geochemical perspectives. (R. E. Taylor, ed.), Noyes

Press, Park Ridge, New Jersey, viii + 360 pp.

, and J. S. E. Carmichael. 1969. The chemical ‘fingerprinting’ of acid volcanic rocks. Calif.

Div. Mines and Geol. Spec. Rept., 100:17-32.

Jackson, T. L. 1974. The economics of obsidian in central California prehistory: application of x-ray
fluorescence spectrography in archaeology. Unpubl. M.A. Thesis, San Francisco, State Univ.,
San Francisco, 216 pp.

Kelley, V. C., and J. L. Soske. 1936. Origin of the Salton volcanic domes, Salton Sea, California. J.
Geol., 44:496—-509.

Muffler, L. J. P., and D. E. White. 1969. Active metamorphism of copper Cenozoic sediments in
the Salton Sea geothermal field and Salton trough, southeastern California. Geol. Soc. Amer.
Bull., 80:157-182.

Nelson, F. W., Jr. 1984. X-ray fluorescence analysis of some western North American obsidians.
Pp. 27-62 in Obsidian studies in the Great Basin. (R. E. Hughes, ed.), Contrib. Univ. California
Archaeol. Res. Facil. 45.

TRACE ELEMENTS OF OBSIDIAN BUTTE 45

Robinson, P. T., W. A. Elders, and L. J. P. Muffler. 1976. Quaternary volcanism in the Salton Sea
geothermal field, Imperial Valley, California. Geol. Soc. Amer. Bull., 87:347-360.

Rogers, A. F. 1934. Salton volcanic domes of Imperial County, California. Geol. Soc. Amer. Proc.
1934:328.

Schamber, F. H. 1977. A modification of the linear least-squares fitting method which provides
continuum suppression. Pp. 241-257 in X-ray fluorescence analysis of environmental samples.
(T. G. Dzubay, ed.), Ann Arbor Science Publishers, Ann Arbor, Mich.

Treganza, A. E. 1942. An archaeological reconnaissance of northeastern Baja California and south-
eastern California. Amer. Antiquity, 8:152-163.

Vane, R. 1977. Trace element sensitivities in geological samples. Nuc. Semicond. Appl. Lab. Rept.
CAL-50, United Scientific Corporation, Mountain View, Calif.

Accepted for publication 9 November 1984.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 46-55
© Southern California Academy of Sciences, 1986

A New Species of Heptacarpus from California, with a
Redescription of Heptacarpus palpator (Owen)
(Caridea: Hippolytidae)

Mary K. Wicksten

Department of Biology, Texas A&M University,
College Station, Texas 77843-3258

Abstract.—A new species of Heptacarpus, a small shrimp marked with brown
dots, is described. It ranges from Santa Rosa Island, California to Guadalupe
Island, Mexico. Related to H. palpator and H. brevirostris, it can be distinguished
from them by having only one large distal spine on the first segment of the
antennular peduncle, and often having a spine at the distal end of the carpus of
the first pereopod. Heptacarpus palpator is redescribed and better distinguished
from H. brevirostris. Heptacarpus palpator ranges from Monterey Bay, California
to the south, while H. brevirostris usually lives north of Monterey Bay.

Members of the family Hippolytidae, often called broken-back shrimp, are
common inhabitants of tidepools, kelp beds, and rocky subtidal bottoms along
the coast of California and northern Mexico. Heptacarpus palpator (Owen) and
H. brevirostris (Dana) have been reported often from shallow areas along the coast
of California (Schmitt 1921; Johnson and Snook 1955; Ricketts et al. 1968; Carlton
and Kuris 1975; Word and Charwat 1976; Chace and Abbott 1980). However,
there are only two keys that distinguish between these two species, that given by
Schmitt (1921) and the one prepared by Word and Charwat (1976). Both distin-
guish between the two species on the basis of the length of the antennal scale to
that of the telson: equal to the telson or shorter in H. brevirostris, or longer than
the telson in H. palpator.

I tried to use the proportions of the antennal scale to the telson in identifying
specimens of Heptacarpus spp. from the coast of California. This proved to be
unsatisfactory. In many specimens labelled H. palpator, the telson was about
0.85 x the length of the antennal scale. To distinguish between the species, careful,
time-consuming measurement with calipers was necessary. Schmitt (1921) noted
the variability in the rostra of the two species, but made no mention of other
parts of the body that might be useful in distinguishing between them.

The original descriptions of H. palpator and H. brevirostris are very short. The
descriptions and illustrations given by Schmitt (1921, as Spirontocaris palpator
and S. brevirostris), and Word and Charwat (1976) also are not extensive. How-
ever, Butler (1980) redescribed H. brevirostris in detail, with a clear illustration.
I was unable to locate the holotype of H. palpator for comparison.

To determine for myself the differences between H. palpator and H. brevirostris,
I examined all the specimens designated as these species in the collections of the
Allan Hancock Foundation (AHF), University of Southern California, totalling
over 200 specimens from 97 stations. I also studied all the specimens of H. palpator

46

A NEW SPECIES OF HEPTACARPUS FROM CALIFORNIA 47

Saree (ceereten
ce

Heptacarpus fuscimaculatus. Ovigerous female, total length in millimeters 14.9. Avalon

Fig. 1.
Harbor, Santa Catalina Island, California, CAS Number 031713.

at the California Academy of Sciences (CAS) as well as specimens labelled H.

brevirostris from south of San Francisco, a total of 32 specimens from 12 stations.
Comparison of all the specimens revealed differences between H. palpator and

H. brevirostris in the rostrum, spines of the carapace, spines of the pereopods,
and ranges in California. However, further examination of the series labelled H.
palpator disclosed an undescribed species. The new species is described herein,

and H. palpator is redescribed.
Heptacarpus fuscimaculatus new species
Figs. 1-2
Rostrum straight, with 1-3 (usually 2) spines on carapace and 2-3 spines on
dorsal margin proper; | or no spines on lower surface. Tip single or bifid. In one
specimen, second spine of carapace bifid. Rostrum slightly exceeding first segment

of antennular peduncle.
Eyes shorter than rostrum, corneas round and pigmented.

Carapace with large antennal, prominent pterygostomial, and no branchiostegal

spines.

Abdominal segments 1-3 rounded at margins. Faint notch of dorsal surface of
second segment. Pleura of fourth segment rounded, each with marginal spine
posterolaterally. Pleuron of fifth segment with prominent posterolateral spine.
Sixth segment about 1.5 x length of fifth segment, with sharp spine on ventrolateral
margin and on lateral margin at articulation with telson. Sixth segment slightly

shorter than telson.

48 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

B

0.5mm

Fig. 2. Heptacarpus fuscimaculatus. A, telson and uropods; B, front in dorsal view; C, dactyl of
third pereopod.

Telson shorter than uropods; with 3—5 pairs dorsolateral spines and 3 pairs
terminal spines with straight setae.

First antenna with prominent, sharp-tipped stylocerite, reaching end of first
segment of its peduncle. Latter with large, curved marginal spine, no spinules.
Second segment about 0.33 x length of first, with sharp terminal spine laterally.
Third segment slightly shorter than second, with sharp terminal spine mesially.
Inner flagellum stout, with about 12 segments and fringe of setae; outer slim, with
at least 15 segments, but often broken during capture.

Second antenna with sharp lower spine on basicerite. Carpocerite about equal
to second segment of antennular peduncle. Scaphocerite shorter than antennular
flagella, 3 x as long as broad; blade shorter than marginal spine, fringed with setae.

Mandible with palp. First maxilla with broad upper endite, small lower endite
and bilobed palp. Second maxilla with small lower endite, bilobed upper endite,

A NEW SPECIES OF HEPTACARPUS FROM CALIFORNIA 49

well developed palp and scaphognathite. First maxilliped with exopod and bilobed
epipod. Second maxilliped with exopod and podobranch. Third maxilliped with
epipod, no exopod, longer than scaphocerite. Ultimate segment setose, ending in
3 sharp claws. Penultimate segment about 0.4 length of ultimate, setose. An-
tepenultimate segment about equal to ultimate, stout, with 2 sharp, stout setae
at articulation with penultimate segment.

First pereopod stoutly chelate, with epipod. Dactyl curved, with tuft of setae
near tip, about equal to palm of chela. Palm of chela about 0.7 x as wide as long.
Carpus about equal to length of palm of chela. Merus longer than palm of chela,
usually with sharp spine or knob near articulation with carpus. Ischium about
0.4x length of merus.

Second pereopod chelate, with epipod. Palm of chela about 2 x length of dactyl.
Carpus with 7 segments, that nearest the propodus the longest. Merus about equal
to 0.5 x carpus, ischium longer than merus.

Third pereopod with epipod. Dactyl stout, about 0.4 x as wide as long, with 4
spines. Tip bifid, claw on flexor margin stouter than that at extreme tip. Propodus
with about 14 spinules, occurring singly or in pairs; 2.6 x length of dactyl. Carpus
short, about 0.5 x length of propodus, broadly overlapping propodus on distal
margin. Merus stout, about 5 x as long as wide, with 2-3 large, sharp distolateral
spines. Ischium shorter than merus, without spines.

Fourth pereopod without epipod, similar to third, but merus 7 as long as
wide and with only 1 or 2 spines. Fifth pereopod without epipod, merus at most
1 spine.

Second pleopods with appendix interna in male, also with appendix masculina.

Uropods longer than telson, fringed with setae. Outer branch with 2 sharp
marginal teeth.

HOLOTYPE: Female, total length in millimeters 12.0. Big Fisherman’s Cove,
Santa Catalina Island, California (33°27'N, 118°28’W), taken by dip net by night
light, among low-growing algae on floating dock, 17 July 1982, water temperature
18°C, Mary K. Wicksten, AHF type number 821.

PARATYPES: California, U.S.A.: Becher’s Bay, Santa Rosa Island (34°0O1'N,
120°02’W), 18 m, sand, 2 Aug. 1938, Velero III sta. 881-38, 1 specimen.—2.5
mi. E. of S. Pt., Santa Rosa Is. (33°53'’45”N, 120°03'40”W-33°53'45’'N,
120°03'40’W), 31-33 m, gravel and red algae, 10 April 1941, Velero ITT sta. 1282-
41, 2 ovigerous females.—E. of Gull Is., S. of Santa Cruz Is. (33°57'15’N,
119°47'15”W-33°57'30’N, 119°48'05”W), 11-18 m, sand and algae, 31 Oct. 1940,
Velero III sta. 1197-40, 10 specimens.—E. of Santa Barbara Island (33°29'N,
119°02'W), shoal to 74 m, sand, 12 Aug. 1938, Velero IIT sta. 895-38, 1 ovigerous
female.—E. of Santa Barbara Is. (33°28’N, 119°00'W), 46-50 m, sand, 27 May
1938, Velero III sta. 975-39, 1 ovigerous female.—E. of Santa Barbara Is., 30
July 1919, Anton Dohrn, 2 specimens.—4 mi. E. of Landing, Santa Barbara Is.
(33°28'40’N, 119°40'00” W-33°28'35’N, 119°30'00”W), 74 m, 28 Aug. 1941, Ve-
lero III sta. 1398-41, 1 ovigerous female.— Big Fisherman’s Cove, Santa Catalina
Island, 17 July 1982, Mary K. Wicksten, 2 specimens other than holotype, private
collection of MK W.—Big Fisherman’s Cove, Santa Catalina Island, among sea-
weed, 29 Sept. 1950, F. Ziesenhenne, 1 specimen.— Big Fisherman’s Cove, Santa
Catalina Island, on dock, 6 Aug. 1984, Mary K. Wicksten, | specimen, Catalina
Mar. Sci. Center reference collection.—Off Howland’s Landing, Santa Catalina

50 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 3. Heptacarpus palpator (Owen). Female, total length in millimeters 44.2. Monterey Municipal
Wharf, Monterey, California; CAS Number 034340.

Island (33°29'28’"N, 118°33'30”N—33°29'45’N, 118°34'10”W), 240-295 m, mud,
10 Sept. 1940, Velero IIT sta. 1183-40, 1 specimen.—1 mi. NW of White Cove,
Santa Catalina Island (33°24’20’N, 118°22'00”W), 4-6 m, among kelp holdfasts,
4 Aug. 1941, Velero III sta. 1378-41, 14 specimens.— 1 mi. ESE of Willow Cove,
Santa Catalina Island (33°22'50’N, 118°19'45”W—33°23'05”N, 118°21'10”W), 63-
92 m, 13 Sept. 1941, Velero III sta. 1403-41, 1 specimen.— White’s Cove, Santa
Catalina Island (33°23'N, 118°22’W), 22-28 m, kelp holdfast, 17 Oct. 1948, Velero
IV sta. 1621-48, 1 ovigerous female.— Avalon, Santa Catalina Island (33°20'45’N,
118°19'45’W), kelp holdfast, 20 Sept. 1938, G. S. Myers, R. L. Bolin and party,
CAS, 1 ovigerous female. —BAJA CALIFORNIA, MEXICO: Melpomene Cove,
Guadalupe Island (28°N, 118°W), 8 Dec. 1946, Carl Hubbs sta. H46-152, 1
specimen.—S. end of Melpomene Cove, Guadalupe Island, 74-83 m, boat Orca,
sta. H50-42, 5 specimens. Except as noted, all specimens are in the collection of
the Allan Hancock Foundation.

Size range. —8.5—15.5 mm in total length.

Remarks. —Heptacarpus fuscimaculatus is related to H. palpator and H. bre-
virostris in having epipods on the first three pereopods and having a rostrum
shorter than the antennular peduncle. All three species have stout, spiny dactyls
on the walking legs, rather than the simple, slender dactyls of H. stimpsoni, a
species which also has epipods on the first three pereopods. Both H. brevirostris
and H. palpator have spinules at the distal end of the first segment of the antennular
peduncle; H. fuscimaculatus has only one spine. Neither of the other two species
has a spine or tubercle at the end of the merus of the first pereopod. Heptacarpus
fuscimaculatus often has three spines on the merus of the third pereopod, H.
palpator usually has two and H. brevirostris often has only one. Heptacarpus
fuscimaculatus usually is found on sandy bottoms or among algae off the islands

A NEW SPECIES OF HEPTACARPUS FROM CALIFORNIA 51

2
Z
—)
s
Z
3
=
ww
ww
—
ws
CT
=

| x

sf

Fig. 4. Heptacarpus palpator (Owen). A, telson and uropods; B, front in dorsal view; C, dactyl of
third pereopod.

of southern California and northwestern Mexico, rather than along the mainland
coast. Heptacarpus fuscimaculatus also is usually smaller as an adult than either
of the other two species.

All of the specimens that I collected at the type-locality, Big Fisherman’s Cove,
were clinging to algae on a floating dock after dark. The grass shrimp Hippolyte
clarki Chace was taken at the same time. In life, three of the animals were trans-
lucent with brown spots formed by chromatophores. The species’ name refers to
these brown spots. The animal taken in 1984 was pale green. A color slide of a

living H. fuscimaculatus has been deposited in the collection of the California
Academy of Sciences.

Heptacarpus palpator (Owen)
Figs. 3-4

Hippolyte palpator Owen 1839:89, pl. 28, fig. 3.
Heptacarpus palpator: Holmes 1900:196, pl. 3, figs. 48, 49.—Holthuis 1947:12.—

52 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Carlton and Kuris 1975:403, pl. 95, figs. 31A, 31B.— Word and Charwat 1976:
122.—Wicksten 1980:205.—Wicksten 1983:26.

Spirontocaris palpator: Rathbun 1904:98.—Schmitt 1921:65, fig. 43.— Ricketts,
Calvin and Hedgpeth 1968:163, fig. 137.

Description. —Rostrum straight, variable in length: reaching at least to cornea,
often to end of first segment of antennular peduncle or slightly beyond; with 2-
3 dorsal spines on carapace and 2-4 dorsal spines on rostrum proper, and 0-2
lower rostral spines; tip usually bifid, sometimes trifid, occasionally single.

Carapace with prominent antennal spine. Pterygostomian spine small, absent
in some individuals.

Abdominal segments 1-3 with rounded pleura. Slight dorsal notch in second
segment. Third segment without dorsal carina. Pleura of segments 4-5 ending in
small, sharp points. Sixth abdominal segment slightly longer than fifth, with 2
sharp lateral points. Telson shorter than uropods, with 4—5 pairs of dorsolateral
spines, the most proximal ahead of middle of telson and set further away from
other spines than the more distal spines are from each other. Apex of telson
pointed, flanked by 5 pair small terminal spines and 1 pair longer spines.

Eyes prominent, cornea pigmented. Eyestalk convex along mesial surface.

First segment of antennular peduncle with 2-3 dorsal spinules and 1 lateral
spine. Stylocerite exceeding first segment. Second segment about 0.3 x length of
first, with sharp lateral spine. Third segment about equal to second, with sharp
dorsal spine. Outer flagellum stout, with dense brush of setae; inner flagellum
whip-like.

Basicerite of second antenna with sharp ventrolateral spine. Carpocerite ex-
ceeding antennular peduncle. Scaphocerite longer than antennular peduncle. Blade
of scaphocerite longer than lateral spine. broader proximally than distally, at most,
about 2 as long as broad.

Mandible with slender incisor process ending in 4 teeth, with two-lobed palp.
Molar process of mandible with numerous spinules. First maxilla with broad
upper endite, small lower endite and bilobed palp. Second maxilla with small
lower endite, bilobed upper endite, well developed palp and scaphognathite. First
maxilliped with exopod and bilobed epipod. Second maxilliped with exopod and
podobranch. Third maxilliped longest appendage of body. Inner branch about
2x length of carpace in male, 1.5 length of carapace in female; with epipod, no
exopod. Ultimate segment witih 6 dark claws at tip, and with straight setae along
its entire length. Penultimate segment about 0.3 x length of ultimate, exceeding
antennular peduncle. Antepenultimate segment about equal to ultimate, with fine
setae.

First pereopod stout, chelate. Fingers of chela about 0.5 x length of palm, tipped
with tufts of setae. Palm 2 as long as wide. Carpus about as long as fingers of
chela. Merus about equal to chela. with lateral row of stout setae. Ischium about
0.3 x merus. Epipod present.

Second pereopod slender, chelate, with epipod. Fingers of chela about 0.4
length of palm, with tufts of setae at tip. Carpus with 7 segments, the first the
longest. Merus 0.6 x length of carpus. Ischium longer than merus.

Third pereopod stout. Dactyl with 2 stout terminal claws and 4 smaller spines,
about 2 as long as wide. Propodus 2.5 length of dactyl, with row of small
spines on flexor margin. Carpus about 0.5 length of propodus, overhanging

A NEW SPECIES OF HEPTACARPUS FROM CALIFORNIA 53

iG

Fig. 5. Heptacarpus brevirostris (Dana). Male, total length in millimeters 42.4. North of Salmon
Creek, Sonoma County, California; AHF sta. 1627-48. Left, dorsal view of front; right, lateral view
of front.

propodus at articulation. Merus about equal to propodus, with usually 2 (rarely
1) sharp ventrolateral spines. Ischium about 0.5 x length of merus.

Fourth and fifth pereopods similar to third, but more slender. One or two spines
on merus of each. No epipods present.

Second pleopod with appendix interna; in male, also appendix masculina.

Uropods fringed with fine setae. Outer branch with 2 sharp lateral spines.

NEOTYPE: The original type specimen of H. palpator came from Monterey,
California. Dr. A. A. Fincham, British Museum (Natural History) informed me
that the type cannot be found. The following specimen therefore is designated as
a neotype: female, total length in millimeters 38.6. Municipal Wharf, Monterey,
Monterey Bay, California (36°37'N, 121°50’W), 8 m, hand net, 2 Feb. 1963, David
Powell, CAS number 035671.

Remarks. — Heptacarpus palpator often has been confused by biologists with H.
brevirostris, a very similar species (Fig. 5). Both are found in intertidal and shallow
subtidal areas on hard bottoms, have rostra that do not reach the end of the second
segment of the antennular penduncle, have stout dactyls on the walking legs with
bifid tips, have similar patterns of spines on the carapace and antennae, and have
epipods on the first three pereopods. To distinguish between the two species, a
combination of features is useful. In H. palpator, the scaphocerite is slightly longer
than the telson; in H. brevirostris, it is equal to or shorter than the telson. In H.
palpator, the spines on the carapace behind the rostrum are not large relative to
those on the rostrum proper; in H. brevirostris, the spines on the carapace form
a descending series toward the rostrum. The spines on the carapace of H. brevi-
rostris are larger than those of the rostrum proper. The rostrum of H. palpator
often has a bifid or trifid tip, and often exceeds the cornea; in H. brevirostris, the
tip usually is single and does not exceed the cornea. The meri of the third—fifth
pereopods of H. palpator usually bear 2, 2, and | spines, respectively; in H. bre-
virostris, these meri bear no more than one spine. Heptacarpus palpator seems to
be slightly smaller than H. brevirostris: the largest specimen recorded had a total
length of 46.6 mm (Schmitt 1921), while Butler (1980) reported H. brevirostris
up to a total length of 62 mm.

54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Heptacarpus palpator has been recorded from San Francisco, California to Bahia
San Gabriel, Isla Espiritu Santo, Gulf of California (Wicksten 1983). Heptacarpus
brevirostris has been reported from Attu, Aleutian Islands to San Francisco (Schmitt
1921; Butler 1980); to south of Carmel, California (Chace and Abbott 1980), and
off Santa Catalina Island and Tanner Bank, California (Word and Charwat 1976).
The northernmost specimens of H. palpator examined by me come from the type
locality and nearby, near Monterey, California. The species occurs commonly in
tidepools in San Luis Obispo County and along the mainland coast of southern
California, from Santa Barbara to San Diego counties. It also occurs along the
outer coast of Baja California, Mexico. It has not been collected often on the
offshore islands of California and Mexico, but has been taken south of San Miguel
Island (Velero III sta. 894-38), at Santa Rosa Island (Velero IIT sta. 995-39), Santa
Cruz Island (sta. H47-89), Farnsworth Bank, off Santa Catalina Island (11-16
Dec. 1970, AHF collections); off Natividad Island, Baja California (Velero IV sta.
1706-49), and between Melpomene Cove and the inner island, Guadalupe Island,
Mexico (Velero IV sta. 1912-49). There is only one record from the Gulf of
California. The species probably rarely ranges south of Magdalena Bay, one of
the most southerly areas of upwelling along the west coast of Baja California
(Dawson 1951).

Heptacarpus brevirostris seems to prefer colder water than H. palpator. It is one
of the most common intertidal shrimps along the coast of British Columbia (Butler
1980). The species has been collected often along the coasts of Washington and
Oregon, U.S.A. I have observed it in tidepools in Humboldt County, California.
It has been taken intertidally and in shallow subtidal waters off Mendocino County,
near Bodega Head in Sonoma County, and along the coasts of San Mateo and
northern Santa Cruz counties, California. I have been unable to examine the
specimens on which Word and Charwat (1976) based their records from southern
California.

Heptacarpus palpator often is found on wharf piles, among algae, and in tide
pools at lower tide levels and in shallow subtidal areas. The deepest record known
to me is 37 m (Santa Monica Bay, California, AHF collections). The color is
variable in life, from translucent with brown lines to a uniform deep brown.

Acknowledgments

I thank Janet Haig, Allan Hancock Foundation, and Dustin Chivers, California
Academy of Sciences, for their help in examining specimens and records, and the
staff of the Catalina Marine Science Center for their hospitality. The illustrations
are by Helen Finney, Texas A&M University. This is contribution number 99 of
the Catalina Marine Science Center.

Literature Cited

Butler, T. H. 1980. Shrimps of the Pacific coast of Canada. Canadian Bulletin of Fisheries and
Aquatic Sciences 202, 280 pp.

Carlton, J. T., and A. M. Kuris. 1975. Keys to decapod Crustacea. Pp. 385—412 in Light’s manual:
Intertidal invertebrates of the central California coast. (R. I. Smith and J. T. Carlton, eds.),
University of California Press, Berkeley.

Chace, F. A., Jr., and D. P. Abbott. 1980. Caridea: the shrimps. Pp. 567—576 in Intertidal inver-
tebrates of California (R. H. Morris, D. P. Abbott and E. C. Haderlie, eds.), Stanford University
Press, Stanford.

A NEW SPECIES OF HEPTACARPUS FROM CALIFORNIA 55

Dawson, E. Y. 1951. A further study of upwelling and associated vegetation along Pacific Baja
California, Mexico. Journal of Marine Research, 10(1):39-58.

Holmes, S. J. 1900. Synopsis of the California stalk-eyed Crustacea. Occasional Papers of the Cal-
ifornia Academy of Sciences, 7:1—262.

Holthuis, L. B. 1947. The Hippolytidae and Rhynchocinetidae collected by the Siboga and Snellius
expeditions with remarks on other species. The Decapoda of the Siboga expedition. Part IX.
Siboga Expeditie 39a®, 100 pp.

Johnson, M. E., and H. J. Snook. 1955. Seashore animals of the Pacific coast. Dover Publications,
Incorporated, New York, 659 pp.

Owen, R. 1839. Crustacea. Pp. 77-92 in The zoology of Captain Beechey’s voyage. H. G. Bohn,
London.

Rathbun, M. J. 1904. Decapod crustaceans of the northwest coast of North America. Harriman
Alaska Expedition, 10:1—210.

Ricketts, E. F., J. Calvin, and J. Hedgpeth. 1968. Between Pacific tides. Stanford University Press,
Stanford, 4th ed, 614 pp.

Schmitt, W. L. 1921. The marine decapod Crustacea of California. University of California Publi-
cations in Zoology, 23:1—470.

Wicksten, M. K. 1980. Crustacea and Pycnogonida. Pp. 196-223 in A taxonomic listing of common

marine invertebrate species from southern California (D. Straughan and R. W. Klink, eds.),

Technical reports of the Allan Hancock Foundation no. 3.

1983. A monograph on the shallow water caridean shrimps from the Gulf of California,
Mexico. Allan Hancock Foundation Monographs in Marine Biology Number 13, 59 pp.
Word, J. Q., and D. Charwat. 1976. Invertebrates of southern California coastal waters. II. Natantia.

Southern California Coastal Water Research Project, El Segundo, California, 238 pp.

Accepted for publication 10 August 1984.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 56-57
© Southern California Academy of Sciences, 1986

Research Notes

Washingtonia robusta Naturalized in Southeastern California

The natural range of the Mexican fan palm, Washingtonia robusta, is limited
to northwestern Mexico (McCurrach 1960) and the Baja Peninsula (Shreve and
Wiggins 1964). However, in California its use as an ornamental is widespread
and the species is the most frequently planted palm in the state (Sunset Editors
1979). It occasionally can be found in disturbed situations immediately adjacent
landscaped areas and David Brown (pers. comm.) reports that it is frequently
encountered along the Colorado River in Arizona and California. To date, it has
not been recorded in undisturbed situations at seeps and springs.

In February 1982 and April 1984, the authors visited a seep in Whipple Wash,
San Bernardino County, California, where a lone palm was found. The specimen
was unburned, approximately 6.5 m in height and had a slender trunk 40 cm in
diameter as measured 1.5 m from the ground. The trunk also flared outward at
its base. The petioles were less than 1.2 m in length, formed a cross-hatched
pattern around the trunk and hung at an angle. These are all characteristics unique
to W. robusta as described by McCurrach (1960), Shreve and Wiggins (1964), and
Wiggins (1980) and we have concluded that the individual does in fact belong to
this species. The palm could have arrived as a seed from the community of Havasu
Palms located 9 km down Whipple Wash at the Colorado River. W. robusta is
an abundant ornamental in this small community and the distance is well within
the dispersal potential of birds or the coyote, Canis latrans (Cornett 1984). Cer-

SOUTHERN CALIFORNIA

Fig. 1. Location of naturalized specimens of Washingtonia robusta at undisturbed sites. The upper
x designates the lone tree in Whipple Wash; the lower x designates the three individuals at Palm Spring
in Anza-Borrego Desert State Park.

56

RESEARCH NOTES 57

cidium floridum, Salazaria mexicana, and Hyptis emoryi were growing within 4
m of the palm indicating the site was undisturbed.

This palm was identified in 1975 as W. filifera by Brown et al. (1976). However,
it probably would not have been of sufficient height at that time to observe the
distinguishing characteristics.

On 3 May 1984 the senior author visited Palm Spring in Anza-Borrego Desert
State Park. Within historic times, this spring had supported several of the desert
fan palms, W. filifera (Edwards 1961). However, these were destroyed by early
settlers and were gone by 1858. In the late 1950’s park rangers attempted to restore
the original plant associates of the spring by planting three palms. It had been
widely assumed that these were palms of the species W. filifera (R. Bloomquist,
pers. comm.). However, it has been concluded that these palms also belong to
the species W. robusta as each of them possesses the characteristics previously
described.

In the past three years we have visited over 95% of the palm oases in California,
Arizona, and Baja California Norte and these are the only locations where we
have found W. robusta occurring under relatively undisturbed conditions above
latitude 32 degrees north.

Acknowledgments

This report was made possible through a grant from the Richard King Mellon
Foundation of Pittsburgh, Pennsylvania.

Literature Cited

Brown, D. E., N. B. Carmony, C. H. Lowe, and R. M. Turner. 1976. A second locality for native
California fan palms (Washingtonia filifera) in Arizona. Journal of the Arizona Academy of
Science, 11:37-41.

Cornett, J. W. 1984. The desert palm oasis. Educational Bulletin #84-1, Desert Protective Council.

Edwards, E.I. 1961. Lost oases along the Carrizo. The Westernlore Press, Los Angeles.

McCurrach, J. C. 1960. Palms of the world. Harper and Brothers, New York.

Shreve, F., and I. L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. Stanford University
Press, Stanford.

Sunset Editors. 1979. New Western Garden Book. Lane Publishing Company, Menlo Park.

Wiggins, I. L. 1980. Flora of Baja California. Stanford University Press, Stanford.

Accepted for publication 7 January 1985.
James W. Cornett,! Jon Stewart,? and Theo Glenn.! '!Natural Science Department,

Palm Springs Desert Museum, Palm Springs, California 92263 and *Living Des-
ert, Palm Desert, California 92261.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 58-61
© Souther California Academy of Sciences, 1986

A Microcephalic Cranium from Catalina Island

In the process of transferring stored skeletal materials from the basement of
the Catalina Island Museum located on the island of Catalina, a microcephalic
skull was discovered among a number of other undocumented skeletal material.
While its exact origins are not known, it is likely that the find was native to either
the Island or adjacent mainland area. Given the relative rarity of microcephaly
in the paleopathological record, the literature is sparse (Guerin 1853; Hrdlicka
1939; Ortner and Putschar 1981) in this area. The skull is virtually complete with
only a few facial bones showing some destruction.

It is difficult to assign an exact biological age to this find because of the distortion
of the usually reliable age-dependent skeletal markers. Arrested growth of the
jaws resulted in the interference of normal tooth eruption which is a prime criterion
for aging a subadult. A reasonable estimate is that the individual was between
seven and nine years of age. Inasmuch as the individual was immature there is
no way to assign a Sex to it.

Seckel (1960) set forth several criteria for defining a true microcephalic. Among
these are a very small cranium, a markedly receding forehead, projecting nasal
bones yielding a “‘bird-headed”’ appearance, widely spaced orbits, underdevel-
opment of the attachment for the neck muscles, a high palate, receding chin etc.

Fig. 1. Lateral View of the Catalina cranium illustrates the strongly receding frontal bone, which
when combined with the elevated nasal structures would produce the “bird-headed”’ appearance. Bar
is 5 cm.

58

RESEARCH NOTES 59

Fig. 2. Comparison with a normal ten year old skull. Note recession of the forehead and the entire
face. Platycephaly, the restricted height of the cranial vault, is readily apparently.

Fig. 3. Fusion of temporal and parietal bones. Bar is 5 cm.

60 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ys ger ie A
aa BO
}
\ /

Fig. 4. Author’s reconstruction of the Catalina cranium.

Employing these criteria, the Catalina cranium clearly demonstrated Seckel’s
syndrome. The circumference of the cranium is 32 cm measured fronto-occipitally
with a cranial capacity of 382 cc employing mustard seed. These measurements
are compatible with an approximately three-month postnatal normal child. The
frontal bone is markedly receding indicating minimal frontal lobe development
(Fig. 1). There is a noticeable keeling in the middle of the forehead. Because of
both reduced cranial capacity and acute frontal angulation, the vault ofthe cranium
is low. Comparison with a normal child approximately ten years old serves to
emphasize the sharply restricted development of the cranial vault (Fig. 2).

Fusion of the cranial bones is limited to a bilateral obliteration of the suture
between the squamosal bone and the parietal bone (Fig. 3). This likely occurred
at an early age. With the exception of the anterior part of the sagittal suture and
the inferior part of the coronal suture all other sutures are open.

Consistent with either diminutive size or nuchal muscle hypoplasia, the occipital
nuchal lines are absent and evidence of strong muscle development is minimal.
Because of the underdevelopment of the mandibular ascending ramus, the external
auditory meatus is relatively low, which, in turn, would have set the ears in a low
position on the head, a feature seen in those with Seckel’s Syndrome.

The elevated and projecting nasal bones and the nasal portion of the maxillary
bones provide the face with what must have been a very prominent nose in life.
In conjunction with the receding forehead, there was likely a well-marked “bird-
headed” appearance (Fig. 4). Given the projecting nasal architecture, the orbits
are thrown into a slightly lateral position. They also appear relatively large but
this is a function of the general hypoplasia of the face (Lim et al. 1973). A relatively

RESEARCH NOTES 61

wide distance between the eyes, hypertelorism, an occasional condition of this
genetic defect is not observed in this specimen.

The palatal portion of the maxillary bone is noticeably arched with the crowded
anterior teeth distorting the dental arch. On both the erupted and unerupted
anterior secondary teeth in both jaws there is moderate to marked absence of
enamel. This is not observed on the primary dentition or what there is left of
them.

The mandible retains both the infantile oblique angle between the horizontal
and vertical portion of the jaw as well as the underdevelopment of these elements
yielding a slightly micrognathic appearance. Chin recession, noted in all instances
of Seckel’s Syndrome, is not a conspicuous feature on the Catalina cranium; when
reconstructed with soft tissue, however the individual would have had a receding
chin.

Given all of the above aberrant osteological features, the Catalina cranium
qualifies as a valid example of the rare Seckel’s Syndrome which Majewski et al.
(1982) believe to be an autosomal recessive condition. Only 17 valid cases of this
disease appear in the clinical record.

Literature Cited

Guerin, J. 1883. Les Microcephales Azteques. Gaz. Med., 42:647-649.

Hrdlicka, A. 1939. Normal, Micro- and Macrocephaly in America. Amer. J. Physical Anthrop., 25:
1-91.

Lim, K. H., and B. Wong, 1973. Ocular anomalies in Seckel’s Syndrome. Austral. and N. Z. J. Med.,
3:520-522.

Majewski, F., and T. Goecke, 1982. Studies of microcephalic primordial dwarfism I: approach to a
delineation of the Seckel Syndrome. Amer. J. of Med. Gen., 12:7-21.

Ortner, D. J., and W. G. Putschar, 1981. Identification of pathological conditions in human skeletal
remains. Smithsonian Contributions to Anthropology, Number 28, 479 pp.

Seckel, H. P., 1960. Bird headed dwarfs. C. C Thomas, Springfield, Ill. 241 pp.

Accepted for publication 20 September 1984.
Stewart Shermis, 9412 Oasis, Westminster, California 92683 or Department of

Anthropology, California State College, 1250 Bellflower Boulevard, Long Beach,
California 90840.

Bull. Southern California Acad. Sci.
85(1), 1986, pp. 62-63
© Southern California Academy of Sciences, 1986

First Record of the Dolphin Steno bredanensis from the
Gulf of California

The cetacean fauna of the Gulf of California, Mexico is quite rich and diverse,
but is poorly documented in the literature (Wells et al. 1981). This account is the
first recorded occurrence of the rough-toothed dolphin (Steno bredanensis) from
this body of water. Records of Steno bredanensis from the tropical and eastern
North Pacific include a few strandings in California, U.S. (Orr 1951; R. Brownell
and C. Woodhouse pers. comm.), the Galapagos Archipelago (Orr 1965), and
incidental catches in tuna purse-seine nets in the eastern tropical Pacific (Perrin
and Walker 1975).

The present specimen was a stranded animal found during an ichthyological
collecting trip in the northern Gulf region. The specimen was located above the
mean high tide mark approximately 1 km south of the town of El Golfo de Santa
Clara, Sonora, Mexico (31°40'N, 114°31’W) on 28 May 1984. The specimen

RO

ay

Se

Fig. 1. The dolphin Steno bredanensis from the Gulf of California.
62

RESEARCH NOTES 63

consisted of a nearly intact rotten carcass which was missing an estimated 15 cm
of the tail including the entire flukes. Judging by the condition of the animal and
the prevailing climate, I estimated the animal had been dead between one and
two weeks. Species identification was based on the following diagnostic characters:
1) in profile, the melon sloped gradually onto the long rostrum (Fig. 1), and 2)
the teeth bore numerous fine vertical ridges (Leatherwood et al. 1982). In addition,
there was a slight hint of lightly pigmented oval and round patches on the body,
especially on the ventral surface, which is typical for adult specimens of Steno.
The following straight line body measurements, in centimeters, were taken : total
length 300 (+15 est.), rostrum to tip of dorsal fin 172, dorsal fin height 19, dorsal
fin length at base 33, rostrum to anterior insertion of flipper 57, anterior insertion
of flipper to flipper tip 47, axilla to flipper tip 31, rostrum to eye 36, and gape 28.

Due to lack of proper permits, the specimen was not collected but photographs
were taken as voucher material (Fig. 1). I wish to thank C. B. Crabtree (UCLA)
for assistance in the data collection and for sponsoring the collecting trip.

Literature Cited

Leatherwood, S., R. R. Reeves, W. F. Perrin, and W. E. Evans. 1982. Whales, dolphins, and porpoises
of the eastern North Pacific and adjacent Arctic waters: A guide to their identification. NOAA
Tech. Rept., NMFS Circ. 444. v + 245 pp.

Orr, R. 1951. Cetacean records from the Pacific coast of North America. Wassman J. Biol., 9:147-

148.

. 1965. The rough-toothed dolphin in the Galapagos Archipelago. J. Mamm., 46:101.

Perrin, W. F., and W. A. Walker. 1975. The rough-toothed porpoise, Steno bredanensis, in the
eastern tropical Pacific. J. Mamm., 56:905-907.

Wells, R. S., B. G. Wursig, and K. Norris. 1981. A survey of the marine mammals of the upper
Gulf of California, Mexico, with an assessment of the status of Phocoena sinus. Nat. Tech. Inf.
Ser. P.B. 81-168792, 51 pp.

Accepted for publication 3 December 1984.

John E. Heyning, Research Associate, Section of Birds and Mammals, Natural
History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles,
California 90007 and Department of Biology, University of California, Los An-
geles.

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
ANNUAL MEETING: MAY 2-3, 1986
The Academy’s 1986 Annual Meeting will be held Friday and Saturday, May 2 and 3, at California
State University, San Bernardino. In conjunction with it, the Southern California Ocean Studies
Consortium and scientists from the Universidad Autonoma de Baja, California Sur, will be presenting
their sixth annual U.S.-Mexico Marine Symposium: and the Desert Studies Consortium of California
State Universities will be presenting a two-day symposium on Desert Ecology. Southern California
Botanists and the American Cetacean Society are also participating in this year’s meeting.

Announcing Memoir #9

BIOLOGY OF THE WHITE SHARK

Papers from a symposium held by the Southern California Academy of Sciences.

Contents include material on shark distribution, ecology, age and growth, visual system,
hematology, cardiac morphology, feeding, temperature, heat production, and exchange,
and attack behavior.

en a er lS ORDER FORM) ae =

Make check or money order payable to Southern California Academy of Sciences, and
mail to:

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

900 Exposition Blvd.

Los Angeles, CA 90007

I would like to order
price of $22.50 per copy.

copies of the “Biology of the White Shark” papers at the

Enclosed is my check for $___________. Please ship my order to me as follows:
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init

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McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp.

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452-458.

Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.),

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CONTENTS

Ecology of the Desert Kit Fox, Vulpes macrotis arsipus, in the Mojave
Desert of Southern California. By Thomas P. O'Farrell and Larry
(FULD ERES OF OPM E EE SIN Sy UI A

A. New Species of Boccardia (Polychaeta: Spionidae) from the Galapagos
Islands and a Redescription of Boccardia basilaria Hartman from
Southern California. By James A. Blake

Notes on Three European Land Mollusks Introduced to California. By
Barry Roth 00) 5 oc) GANA sae AO
The Taxonomic Status of Two North American Lithurge (Hymenoptera:
Megachilidae). By Roy R. Snelling
Trace Element Composition of Obsidian Butte, Imperial County, California.
By Richard E. Hughes
A New Species of Heptacarpus from California, with a Redescription of
Heptacarpus palpator (Owen) (Caridea: Hippolytidae). By Mary K.
Wicksten

Research Notes

_-Washingtonia robusta Naturalized in Southeastern California. By James W. Cornett, Jon

Stewart ana: ENeo Glertry oa ei ye pa ORIN LE

A Microcephalic Cranium from Catalina Island. By Stewart Shermis 0

First Record of the Dolphin Steno bredanensis from the Gulf of California. By John E.
VAC Ying oes Le NWN OIA UG) ae ee ee ARENT S aa

16

2D

as)

315

46

COVER: Desert fan palms, Washingtonia filifera, at Mopah Springs in the Turtle Mountains, San

Bernardino County, California. Photograph by James W. Cornett, Palm Springs Desert

Museum.