Occasional papers of the California Academy of Sciences

Edward L. Kessel, Editor

OCCASIONAL PAPERS
OF THE

California Academy of Sciences

No. 108, 15 pages, 28 figures

PHALANGODIDAE FROM CAVES

IN THE SIERRA NEVADA (CALIFORNIA)

WITH A REDESCRIPTION OF THE TYPE GENUS

(Opiliones:Phalangodidae)

By

Thomas S. Briggs

U

\ 2 1974

Woods Hole, Mass.

SAN FRANCISCO
PUBLISHED BY THE ACADEMY
March 14, 1974

^

OCCASIONAL PAPERS

OF THE

California Academy of Sciences

No. 108, 15 pages, 28 figures.

PHALANGODIDAE FROM CAVES

IN THE SIERRA NEVADA (CALIFORNIA)

WITH A REDESCRIPTION OF THE TYPE GENUS

(Opiliones:Phalangodidae)

By

Thomas S. Briggs

Research Associate, California Academy of Sciences
and Galileo High School, San Francisco, California

Many of the small limestone caves in the Mother Lode
district of the Sierra Nevada possess a varied biota
which includes a distinctive genus of laniatorid
harvestmen. This group has previously been known only by
one species, Banksula calif ornioa (Banks) which was
collected in Alabaster Cave, El Dorado County, California
at the turn of the century." Several new species in the
genus Banksula were collected during an investigation of
Sierra Nevada caves with the San Francisco Bay Chapter of
the National Speleological Society.

It is now apparent that the genus Banksula is entirely
confined to caves even though all but one species possess
eyes. The only laniatorid opilionids found on the ground
in the Mother Lode district are in an unrelated genus of
Phalangodidae. This surface-inhabiting genus does include
one cavernicolous example in the Sierra Nevada {Sitaloina
oloughensis Briggs, 1967), but it is found south of the
Mother Lode district in a different limestone formation.
In spite of broad separation of deposits of Mother Lode
(Calaveras) limestone by deep river valleys the allopatric
populations of Banksula are morphologically similar. This
phenomenon of a regional population of allopatric cave

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

laniatorids being related to each other but unrelated to the
regional population of surface laniatorids exists in several
parts of the United States and elsewhere and will be
discussed in a future paper.

The Mother Lode caves are found in small limestone
outcrops and are unprotected from various destructive
types of human use. The type species, Banksula calif ovnioa ,
is only known from Alabaster Cave. From the time
Alabaster Cave was partially destroyed by quarrying no
further specimens have been found, and the species may be
extinct. Another cave, McLean's Cave, is important because
among other organisms it contains sympatric species of
Banksula. This cave will be flooded by a new dam on the
Stanislaus River. Several caves, particularly Moaning Cave,
are threatened by "improvements" for recreational use.

Several authors have placed Banksula calif ovnioa in
the genus Phalangodes . The genus Phalangodes has long been
the recipient of a variety of unrelated species from Europe,
United States, and the Carribean. The type species,
Phalangodes avmata Tellkampf, occurs in the Mammoth Caves
of the state of Kentucky. An examination of this and
related species indicates that Phalangodes is restricted
to the southeastern United States, possibly limited to
caves. To help clarify the morphology of both this genus
and Phalangodidae as a whole, redescription of Phalangodes
will be given based on specimens from Mammoth Cave and
nearby Bypass Cave.

During this study it was found desirable to expand the
folded velum of the opilionid penis. This was accomplished
by brief treatment with 10 percent KOH as is done with the
palpi of male spiders. Such a procedure is necessary for
a complete description of the genitalia of most male
harvestmen in the superfamily Gonyleptoidea (Oncopodoidea) .

All types collected in this study are deposited in the
collections of the California Academy of Sciences.

PHALANGODES Tellkampf, 1844

Fused segments of scute not delineated by grooves , first
segment without a median line. Eye tubercle small, unarmed,
clearly separated from anterior margin of scute. Labial
processes visible between, but not anterior to, endites of
second coxae. Spiracles visible. Sternum narrow, broaden-
ing beneath operculum. Tergits reduced, lateral sclerites
present. Tarsi of third and fourth legs without scopulae
and with simple untoothed double claws. Disti tarsi of first
leg with two segments, of second with three. Tarsi of first
legs with four or five segments. Palpal tarsi with four
major lateral spines, femur without dorsal spines. Male
with ventral projections on endites of second coxae. Penis
with apical process (dorsal plate?) on narrow, unhinged
aedeagus ; ventral plate bifurcate. Juvenile with large
aroleum on hind claws.

TYPE SPECIES. Phalangodes avmata Tellkampf (see
figs. 1-7) .

No. 10 8] BRIGGS: PHALANGODIDAE

SPECIMENS EXAMINED. Kentucky: Mammoth Cave, 3 January
19 31, H. Dietrich; Bypass Cave, near Bowling Green, Warren
County, 31 August 1967, S. Peck and A. Fiske.

BANKSULA Roewer, 19 49

Fused segments of scute weakly delineated by grooves,
first segment without a medial line. Eye tubercle of
normal size, unarmed, clearly separated from anterior
margin of scute. Labial processes project anterior to
endites of second coxae. Spiracles visible. Sternum
narrow, broadening beneath operculum. Tergites of normal
size, lateral sclerites present. Tarsi of third and fourth
legs without scopulae and with simple untoothed double
claws. Disti tarsi of first leg with two segments, of
second with three. Tarsi of first legs with four segments.
Palpal tarsi with four major lateral spines, femur with
six to eight dorsal spines. Males without ventral pro-
jections on endites. Penis without apical process on
aedeagus, hinged aedeagus and dorsal plate expand above
ventral plate, dorsal and ventral plates bifurcate. Juve-
niles with large aroleum on hind claws.

TYPE SPECIES. Scotolemon oalifornicus Banks.

Key to the Males of Banksula

1. Operculum small, posterior margin continuous with

posterior margin of fourth coxae; apex of

aedeagus not enclosed in sheath 2

Operculum large, posterior margin behind posterior
margin of fourth coxae; apex of aedeagus enclosed
in sheath 4

2 . Without corneas and retinae

Banksula calif ovnioa (Banks)

With corneas and part or all of retinae 3

3. Second endites with a concave ectal margin of

constant radius Banksula galilei , new species

Second endites with an angular invagination of
the ectal margin, interior apex close to
posterior margin Banksula tuolumne , new species

4. Aedeagus and enclosing sheath held within

bifurcate dorsal plate... Banksula g rah ami , new species
Aedeagus and enclosing sheath folds anterior

to recurved prongs of dorsal plate

Banksula melones , new species

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Banksula oalifornioa (Banks).
(Figures 8-11. )

Sootolemon oalifornious Banks, 1900, J. N. Y. Entomol. Soc. ,

vol. 10, p. 200; 1901, Amer. Natur. , vol. 35, p. 672;

1904, Proc. Calif. Acad. Sci., ser. 3, vol. 3, p. 363.
Phalangodes oalifornioa , Roewer, 1912, Arch. Naturgesch. ,

vol. 78A, fasc. 3, p. 142; 1923, Die Weberknechte der

Erde , p. 105.
Banksula oalifornioa , Roewer, 1949, Senckenbergiana, vol.

30, p. 33.

LECTOTYPE. Female. Body length 2.06mm. Scute length
1.25mm. Scute width 1.3 3mm. Eye tubercle length 0.21mm.
Eye tubercle width 0.38mm.

II

III

IV

Palpus

Trochanter

0 . 1mm.

0. 2mm.

0. 2mm.

0. 2mm.

0 . 2mm

Femur

1.1

1.4

1.1

1.4

0.7

Patella

0.3

0.4

0.3

0.4

0.4

Tibia

0.8

1.3

0.9

1.3

0.6

Metatarsus

0.8

1.0

1.0

1.3

-

Tarsus

0.8

1.5

0.8

1.1

0.8

Total

3 . 9mm.

5 . 8mm.

4. 3mm.

5. 7mm.

2. 7mm

Scute with cheliceral sockets, not tuberculate, segmen-
tation not apparent. Eye tubercle subconical, tuberculate,
retinae absent, corneas very small or absent. Tergites
with row of tubercles at posterior margins. Second endites
large, setose, project anteriorly as triangular lobes.
Labial processes project anterior to second endites.
Operculum small, subtriangular , posterior margin adjacent
to mesal apex of posterior margin of fourth coxae.

Chelicerae setose, basal segments with ectal spur.

Palpi with six major spines on dorsum of femora.

Tarsal formula 4-6-5-6.

Color of body concolorous yellow.

PARALECTOTYPE . Male. Body length 1.8 8mm. Scute
length 1.41mm. Scute width 1.36mm. Eye tubercle length
0.31mm. Eye tubercle width 0.38mm.

II

III

Trochanter

0. 2mm.

0. 2mm.

0. 2mm.

Femur

1.1

1.6

1.3

Patella

0.4

0.5

0.3

Tibia

-

0.4

1.0

Metatarsus

-

-

1.1

Tarsus

-

-

0.9

Total

-

-

4. 8mm.

IV

0. 2mm.

1.4

0.4

1.4
1.4
0.9

5 . 7mm.

Palpus

0 . 3mm.
0.9
0.6
0.7

0.7

3 . 2mm.

No. 10 8]

BRIGGS: PHALANGODIDAE

Similar to female but with narrower operculum and more
robust palpal femur. Penis with bifurcate ventral plate
bearing setae on recurved distal processes. Articulated
dorsal plate retracted anterad, bifurcate with acute distal
processes. Base of dorsal plate smooth. Aedeagus un-
sheathed at apex, unfolds anterior to dorsal plate when
turgid.

TYPE DESIGNATIONS. Lectotype female, paralectotype
male and paralectotype females from the Nathan Banks
type series in the collection of the Harvard Museum of
Comparative Zoology. Types are from Alabaster Cave, El
Dorado County, California.

REMARKS. A thorough search has failed to locate speci-
mens in the remainder of Alabaster Cave. The male para-
lectotype does not have complete first and second legs.

Banksula tuolumne Briggs, new species.
(Figures 12-17.)

HOLOTYPE. Male. Body length 1.80mm. Scute length
1.33mm. Scute width 1.36mm. Eye tubercle length 0.2 8mm.
Eye tubercle width 0.39mm.

II

III

IV

Palpus

Trochanter

0. 2mm.

0 . 2mm.

0 . 2mm .

0. 2mm.

0. 2mm.

Femur

1.0

1.4

1.2

1.1

0.9

Patella

0.4

0.5

0.3

0.5

0.5

Tibia

0.9

1.4

1.1

1.5

0.6

Metatarsus

1.0

1.3

1.2

1.5

-

Tarsus

0.9

1.7

1.0

1.3

0.8

Total

4. 4mm.

6 . 5mm.

5. 0mm.

6 . 1mm.

3. 0mm.

Scute with cheliceral sockets, moderately tuberculate,
segmentation delineated by rows of tubercles. Eye tubercle
subconical, tuberculate, with small corneas and depigmented
retinae. Tergites with row of tubercles. Second endites
large, setose, narrowest behind midpoint at deep angle.
Spatulate labial processes project anterior to second
endites. Operculum small, subtriangular , posterior margin
adjacent to mesal apex of posterior margin of fourth coxae.

Chelicerae setose, basal segments with ectal spur.

Palpi with eight major spines on dorsum of femora.

Tarsal formula 4-6-5-6.

Color of body concolorous orange-yellow.

Penis with bifurcate ventral plate bearing setae on
recurved distal processes. Articulated dorsal plate bi-
furcate with acute distal processes arched mesodorsad.
Dorsal plate retracted posterad. Base of dorsal plate
smooth. Aedeagus unsheathed at apex, unfolds anterior to
dorsal plate when turgid.

6

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

ALLOTYPE. Female. Similar to male.

TYPE SPECIMENS. Holotype male and allotype female:
under wood in lower level, Tuolumne Crystal Cave, near
Tuolumne, Tuolumne County, California, 22 February 1969,
T. Briggs. Nine paratypes : same locality, 22 February
1969, T. Briggs.

Banksula galilei Briggs, new species.
(Figures 18-22.)

HOLOTYPE. Male. Body length 1.62mm. Scute length
1.12mm. Scute width 1.12mm. Eye tubercle length 0.20mm.
Eye tubercle width 0.29mm.

II

III

IV

Palpus

Trochanter

0. 2mm.

0 . 2mm.

0. 2mm.

0 . 2mm.

0 . 2mm.

Femur

1.0

1.1

1.0

1.1

0.7

Patella

0.3

0.4

0.3

0.4

0.5

Tibia

0.7

1.1

0.9

1.1

0.5

Metatarsus

0.8

0.9

1.0

1.1

-

Tarsus

0.8

1.2

0.8

1.0

0.3

Total

3 . 8mm.

4. 9mm.

4. 2mm.

4. 9mm.

2 . 2mm.

Scute with cheliceral sockets, lightly tuberculate ,
segmentation delineated by rows of small tubercles. Eye
tubercle subconical, tuberculate, with small corneas and
retinae. Tergites with row of tubercles. Second endites
large, setose, narrowest at midpoint, ectal margin concave.
Spatulate labial processes project anterior to second
endites. Operculum small, apically rounded, posterior
margin adjacent to mesal apex of posterior margin of fourth
coxae .

Chelicerae setose, basal segments with ectal spur.

Palpi with seven major spines on dorsum of femora.

Tarsal formula 4-6-5-6.

Color of body concolorous orange-yellow.

Penis with bifurcate ventral plate bearing setae on
recurved distal processes. Articulated dorsal plate bi-
furcate with acute distal prongs arched dorsad. Basal stem
of dorsal plate smooth. Aedeagus unsheathed at apex, un-
folds anterior to dorsal plate when turgid.

ALLOTYPE,
all size.

Female. Similar to male but larger in over-

TYPE SPECIMENS. Holotype male: under breakdown in pit
room, Lime Rock Caves, near Auburn, Placer County, Cali-
fornia, 21 December 1966, V. F. Lee and T. Briggs. Allotype
female: same locality, 2 June 1966, A. Jung, K. Horn, and
T. Briggs. Five paratypes: same locality, 21 December and
2 June 19 66, A. Jung, K. Horn, V. F. Lee, and T. Briggs. Two
paratypes: Lime Rock Cave #3, same locality, 2 June 1966,
K. Horn and T. Briggs.

No. 10 8] BRIGGS: PHALANGODIDAE

Banksula grahami Briggs , new species.
(Figures 23-26.)

HOLOTYPE. Male. Total body length 1.6 3mm. Scute
length 1.2 0mm. Scute width 1.2 3mm. Length of eye tubercle
0.22mm. Width of eye tubercle 0.30mm.

I II III IV Palpus

Trochanter

0 . 2mm.

0 . 2mm.

0. 2mm.

0 . 2mm.

0 . 2mm

Femur

0.9

1.3

1.1

1.3

0.7

Patella

0.4

0.4

0.3

0.4

0.4

Tibia

0.7

1.1

0.8

1.2

0.6

Metatarsus

0.7

1.0

1.0

1.2

—

Tarsus

0.8

1.7

0.8

1.0

0.7

Total

3 . 7mm.

5. 7mm.

4. 2mm.

5 . 3mm.

2 . 6mm

Scute with cheliceral sockets, segmentation delineated
by small tubercles. Eye tubercle subconical, slightly
tuberculate, with small corneas and retinae. Tergites with
row of small tubercles. Second endites setose, mesally
broad and rounded. Labial processes spatulate, project
anterior to second endites. Operculum large, posterior
margin posterior to mesocaudal apex of hind coxae.

Chelicerae with ectal spur on basal segments.

Palpi with six major spines on dorsum of femora.

Tarsal formula 4-6-5-6.

Color of body concolorous yellow-orange.

Penis with bifurcate ventral plate bearing setae on
recurved distal processes. Articulated dorsal plate
bifurcate, base broad and slightly wrinkled. Apex of
aedeagus enclosed in rounded sheath and held within
distal portion of dorsal plate.

ALLOTYPE. Female. Similar to male. Operculum
slightly thickened along anterior margin.

TYPE SPECIMENS. Holotype male and allotype female:
under wood and rocks, Moaning Cave, near Vallecitos,
Calaveras County, California, 22 December 1968, T. Briggs.
Two paratypes: Moaning Cave, (56°F.), near Vallecitos,
Calaveras County, California, 22 August 1963, R. E.
Graham. Two paratypes: same locality, 22 December 196 8,
T. Briggs.

OTHER LOCALITIES. Tuolumne County: under rocks,
Crystal Palace Cave, near Columbia, 4 November 1967, V. F.
Lee and T. Briggs; under talus in main passage, McLean's
Cave, near Columbia, 13 May 196 7, T. Briggs; same locality,
24 June 196 7, K. Horn and T. Briggs.

REMARKS. Some specimens have as many as eight dorsal
spines on the palpal femur. Seven segments are found on
the second or fourth tarsus of some individuals.

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

Banksula melones Briggs, new species.
(Figures 27-28.)

HOLOTYPE. Male. Body length 2.22mm. Scute length
1.33mm. Scute width 1.40mm. Eye tubercle length 0.39mm.
Eye tubercle width 0.41mm.

II

III

Trochanter

0 . 2mm.

0 . 2mm.

0 . 2mm

Femur

1.1

1.5

1.1

Patella

0.4

0.5

0.3

Tibia

0. 8

1.3

1.0

Metatarsus

0.9

1.1

1.1

Tarsus

0.9

1.7

0.9

Total

4 . 3mm.

6 . 3mm.

4 . 6mm

IV

0 . 3mm.

1.3

0.5

1. 3

1.3

1.1

5 . 8mm.

Palpus

0 . 2mm.
0.9
0.5
0.6

0.8
3. 0mm.

Scute with cheliceral sockets, segmentation delineated
by small tubercles. Eye tubercle subconical, slightly
tuberculate ; with large, well developed eyes. Tergites with
row marginal tubercles. Second endites setose, sub-
triangular. Labial processes project distoventrally from
second endites. Operculum large, posterior margin pos-
terior to mesocaudal apex of hind coxae.

Chelicerae with ectal spur on basal segments.

Palpi robust, eight major spines on dorsum of femora.

Tarsal formula 4-6-5-6.

Color of body concolorous yellow-orange.

Penis with shallow bifurcation of ventral plate, ectal
setae present on recurved distal processes. Articulated
dorsal plate bifurcate, base broad and corrugated. Apex of
aedeagus in truncate sheath which unfolds anterior to
acute, recurved distal processes of dorsal plate.

ALLOTYPE

Female. Similar to male

TYPE SPECIMENS. Holotype male and allotype female:
middle level, Quarry Cave, near Columbia, Tuolumne County,
California, 24 May 1969, G. Leong, W. Rauscher, and T.
Briggs.

OTHER LOCALITIES. Tuolumne County: under talus in
main passage, McLean's Cave, near Columbia, 13 May 1967,
V. F. Lee and T. Briggs; same locality, 24 June 1967,
K. Horn and T. Briggs; same locality, 17 June 19 67, V. F.
Lee and T. Briggs.

No. 10 8] BRIGGS: PHALANGODIDAE 9

ACKNOWLEDGMENTS

The author wishes to thank Herbert W. Levi of the
Harvard Museum of Comparative Zoology for the loan of the
type of Banksula californiaa, Willis J. Gertsch, John A. L,
Cooke, and Stewart B. Peck for the loan of specimens of
Phalangodes avmata and Richard E. Graham for specimens
of Banksula from Moaning Cave. Drawings were made by Jack
Tom and Kevin Horn. Much of the field investigation was
done by Kevin Horn, Albert Jung and Vincent F. Lee while
they were science students at Galileo High School.

10

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

FIGURES 1-7. Phalangodes armata Tellkampf. FIGURE 1.
Lateral view of female. FIGURES 2-3. Claws of hind tarsus
FIGURE 4. Ovipositor. FIGURE 5. Dorsal view of female.
FIGURES 6-7. Lateral and ventral views of penis.

No. 108]

BRIGGS: PHALANGODIDAE

11

10

«■#<»

FIGURES 8-11. Banksula oalifovnica (Banks). FIGURE 8.
Dorsal view of lectotype. FIGURES 9-10. Lateral and ventral
views of penis. FIGURE 11. Lateral view of lectotype.

12

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

FIGURES 12-17. Banksula tuolumne Briggs , new species.

FIGURE 12. Dorsal view of male. FIGURES 13-14. Claws of

hind tarsus. FIGURES 15-16. Ventral and lateral views of
penis. FIGURE 17. Lateral view of male.

No. 108]

BRIGGS: PHALANGODIDAE

13

FIGURES 18-22. Banksula galilei Briggs, new species.
FIGURE 18. Ventral view of penis. FIGURE 19. Claws of
hind tarsus. FIGURE 20. Dorsal view of male. FIGURE 21,
Ovipositor. FIGURE 22. Lateral view of male.

14

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

24

25

FIGURES 23-26. Banksula grahami Briggs , new species.
FIGURE 23. Lateral view of male. FIGURES 24-25. Lateral
and ventral views of penis. FIGURE 26. Dorsal view of male

No. 108]

BRIGGS: PHALANGODIDAE

15

27

28

FIGURES 27-28. Lateral and ventral views of the penis
of Banksula melones Briggs, new species.

MBL/WHOI LIBRARY

UH 1TFZ N