A new species and range extensions for three other species of pebblesnails (Lithoglyphidae, Fluminicola) from the upper Klamath basin, California-Oregon
Author
Liu, Hsiu-Ping
Author
Hershler, Robert
text
ZooKeys
2019
812
47
67
http://dx.doi.org/10.3897/zookeys.812.29205
journal article
http://dx.doi.org/10.3897/zookeys.812.29205
1313-2970-812-47
940784938803416F83DB04B0BEACBD92
Fluminicola klamathensis Liu & Hershler
sp. n.
Figures 5, 6, 7, 8A
Types.
Holotype, USNM 1144499 (a cleaned shell), Lost River at Stukel Bridge, Klamath County, Oregon,
42.0825N
,
121.6617W
, 10/5/1997, Terrence J. Frest and Edward J. Johannes. Paratypes, USNM 1468970 (a large series of dry shells and alcohol-preserved specimens), from same lot.
Referred material.
OREGON. Klamath County: USNM 1207966, Lost River at Stukel Bridge, USNM 1144894, USNM 1469075, USNM 1469082, USNM 1469090, Wood River, south spring source (
42.7372N
,
121.9775W
), USNM 1469072, USNM 1469077, USNM 1469078, USNM 1469080, Tecumseh Spring (
42.6424N
,
121.9432W
), USNM 1144346, USNM 1190088, Camporee Spring (
42.4308N
,
122.0614W
), USNM 883517, USNM 1144348, USNM 1190089, USNM 1207965, USNM 1225874, Harriman Spring, outflow of main spring (
42.4673N
,
122.1009W
), USNM 1469076, USNM 1469086, Big Springs at Bonanza (
42.1982N
,
121.4004W
), USNM 1469074, USNM 1469088, Lost River, below Harpold Dam (
42.1702N
,
121.4530W
).
Diagnosis.
A large
Fluminicola
(shell height, 6.5-8.4 mm) with a subglobose to ovate-conic shell often having an eroded spire.
Fluminicola klamathensis
is readily distinguished by its penis, which does not gently taper along its length as in other congeners, but instead abruptly narrows distally and has a well demarcated, short filament. This new species is further differentiated from closely similar and frequently sympatric
F. modoci
in its generally darker colored shell periostracum, broad central cusps on the central and lateral radula teeth, very short outer wing of the lateral teeth, small number of cusps on the inner and outer marginal teeth, light pigment on the dorsal surface of the penis, large core of internal dark pigment in the distal section of the penis, and the fairly large seminal receptacle.
Description.
Shell (Fig. 5
A-F
) subglobose to ovate-conic, spire often eroded in large part, whorls (in specimens having a complete spire), 4.25-4.5. Teleoconch whorls low to medium convex, sometimes weakly shouldered. Aperture pyriform; inner lip complete, variably thickened, broadly adnate to parietal wall; columellar shelf narrow or extending over much of umbilical region. Outer lip thin, prosocline. Umbilicus absent or a narrow slit, umbilical region sometimes excavated. Shell white, periostracum brown, fairly thick, sometimes covered with black deposits. Shell measurements and whorl count data are summarized in Table 2.
Figure 5. Shells of
F. klamathensis
, sp. n. A Holotype, USNM 144499 B, C USNM 1469076 D USNM 1469078 E USNM 1144346 F USNM 1144894. Scale bar: 1.0 mm.
Table 2. Shell parameters for
F. klamathensis
. Measurements are in mm.
Operculum (Fig. 6
A-C
) as for genus; muscle attachment margin little thickened on inner side; rim sometimes present on inner side near outer edge (Fig. 6B). Radula (Fig. 6
D-H
) as for genus; dorsal edge of central teeth concave, lateral cusps 2-5, hoe-shaped; basal cusp 1-5. Lateral teeth having 2-4 cusps on inner side and 3-5 cusps on outer side; outer wing slightly longer than length of cutting edge. Inner marginal teeth with 11-19 cusps, outer marginal teeth with 11-22 cusps. Radula data are from USNM 144346, USNM 1468970.
Figure 6. Opercula and radula,
F. klamathensis
, sp. n. A, B Opercula (outer, inner sides), USNM 1468970 C Operculum (inner side), USNM 144348 D Portion of radular ribbon, USNM 1468970 E, F Central teeth, USNM 144346, USNM 1144348 G, H Lateral teeth, USNM 1144346, USNM 1144348. Scale bars: 500
µm
(
A-C
); 20
µm
(
D-H
).
Snout
, cephalic tentacles grey or black, pigment light around eyespots; pallial roof, visceral coil usually light brown; foot variably pigmented dorsally, sole pale. Ctenidial filaments 33-36 (N = 5), broadly triangular. Glandular oviduct and associated structures shown in Figure 7A, B. Coiled oviduct circular, anterior arm kinked, posterior arm sometimes having small accessory pouches containing sperm. Bursa copulatrix large, ovate or globular, partly overlapped by albumen gland. Bursal duct narrow, much shorter than bursa. Seminal receptacle medium-sized, pouch-like, partly overlapped by albumen gland. Albumen gland having small pallial component. Capsule gland slightly shorter than albumen gland. Genital aperture a small, terminal pore. Penis (Figs 7C, D, 8A) large, base rectangular, often having a distinct, lobe-like swelling along inner edge distally (Fig. 7D); distal end of penis blunt, with short, narrow, filament. Distal section of penis having dense core of internal black pigment; dorsal surface pale or lightly pigmented. Penial duct near inner edge, narrow, nearly straight.
Figure 7. Reproductive anatomy,
F. klamathensis
sp. n. A Female glandular oviduct and associated structures (viewed from left side), USNM 1225874 B Bursa copulatrix and seminal receptacle, USNM 1225874 C, D Penis (dorsal surface), USNM 1225874, USNM 1469090. Ag albumen gland Bu bursa copulatrix Cg capsule gland Cov coiled oviduct Dsr seminal receptacle duct Ga genital aperture In inner edge of penis Ou outer edge of penis Pd penial duct Pf penial filament Pw posterior wall of pallial cavity Sr seminal receptacle Vc ventral channel of capsule gland. Scale bars: 1.0 mm (
A-C
); 500
µm
(D).
Etymology
.
The species name is an adjectival geographic epithet referring to the distribution of this pebblesnail in the (upper) Klamath River basin.
Distribution.
Large, spring-influenced habitats in the UKL.
Remarks.
We selected the Lost River at Stukel Bridge as the type locality because the pebblesnails in this population do not have apically eroded shells. However, we did not have suitably relaxed material from this locality for anatomical study and thus used specimens from Harriman Springs for this purpose.
The shells of
F. klamathensis
and
F. modoci
can be difficult to distinguish although the former usually has darker, thicker periostracum, and tends to be larger when found in sympatry. As noted above, these two species are most readily differentiated by the shape and pigmentation of the penis (Fig. 8).
Figure 8. Photographs of ethanol preserved, relaxed specimens of
F. klamathensis
sp. n. (A) and
F. modoci
(B) from the Wood River south headspring, showing the differences in penial morphology A USNM 1469072 B USNM 1144736. Pn penis. Scale bar: 200
µm
.
Sixteen (16) COI and 13 cytB haplotypes were detected in
F. klamathensis
(Suppl. materials 2, 3, respectively).
The "Tall pebblesnail" (also referred to as
Fluminicola
n. sp. 2) that was recognized by Frest and Johannes in their UKL contract reports (also see
Frest and Johannes 1999
) and subsequently included in the Northwest Forest Plan as a Survey and Manage species, corresponds to
F. klamathensis
.