Amplaria muiri, Shear, William A. & Krejca, Jean K., 2007

Shear, William A. & Krejca, Jean K., 2007, Revalidation of the milliped genus Amplaria Chamberlin 1941 (Diplopoda, Chordeumatida, Striariidae), and description of two new species from caves in Sequoia and Kings Canyon National Parks, California, Zootaxa 1532, pp. 23-39 : 28-35

Amplaria muiri , n. sp.

Figs. 4–15 , 24, 26 , 27

Types: Male holotype and female and male paratypes from Crystal Cave, Sequoia National Park, Tulare Co., California, collected 15 July 2003 by Jean Krejca, V. Loftin and S. Fryer, deposited in Field Museum of Natural History, Chicago (FMNH).

Diagnosis: Distinct from the nearby A. adamsi , n. sp., in the larger size, fewer ocelli, paler pigmentation, and details of the gonopods as illustrated.

Etymology: After John Muir, famous naturalist of the late nineteeth and early twentieth century, whose name is forever connected with the Sierra Nevada, which he called “the Range of Light.”

Description: Male. Length, 16.5 mm, width, 1.2 mm. Three, four or five small, irregularly shaped black ocelli in one or two rows. Crest 6 (lateralmost) of collum incomplete, obscure; more prominent on metazonites 2, 3, complete on metazonite 4. Segmental setae of collum to metazonite 5 aciculate, twice as long as intercrest distance, becoming shorter from metazonite 6 posterior, blunt, subclavate, as long as intercrest distance or shorter. Epiproct broad, spatulate. Color of most specimens yellowish white ( Figs. 24, 26 , 27 ), few have slight reticulate pattern of pale purplish brown on posterior margins of metazonites 24–29 and epiproct; legs and antennae white. Secondary sexual modifications as described for genus.

Gonopods ( Figs. 7–11 ) robust, anterior angiocoxites appressed in midline, apically with three unequal teeth, deltoid lateral branch sharply, evenly curved; posterior angiocoxites with flagellar sheathing branch stout, apically flaring, posterior branch inflated, with curved, toothed ridge. Colpocoxites typical of genus (note: poorly sclerotized lateral lobe of colpocoxite collapses in SEM preparation). Legpair 9 as in Fig. 15 .

Female: length, 15.2 mm, width, 1.0 mm. Except for secondary sexual characters, much as male.

to be contineud.

Distribution; See Table 1 and fig. 23.

Habitat, abundance and life history: Eighty-four specimens from Bear Den Cave, Carmoe Crevice, Crystal Cave, Hurricane Crawl Cave, Lange Cave, and Pet Cemetery Cave were positively identified, and another nine immature and female specimens from Lilburn Cave and Weisraum Cave were tentatively identified ( Table 1 ), and the following data are based on those 93 individuals. Air temperatures were recorded for 75 of those collections and the average air temperature where specimens were collected was 10.4 deg C (range = 8.3 to 12.5 deg C). Substrates were recorded for 37 of those collections: 43% (16/37) were found on a silt floor, 24% (9/37) were found on a dead small mammal, 16% (6/37) were found on roots, 11% (4/37) were found on calcite, and 5% (2/37) were found under a rock. Of those 37 collections, 73% (27/37) were associated with some kind of energy source, including fungus, roots, a dead mammal or bait on a pitfall trap. For all 93 specimens, the average distance they were found into the cave was 80 m (range = 0 to 166 m). Also on average one A. muiri was found per 7 linear meters of cave passage (range = 1 m to 48 m). For 82 specimens with records of search effort, on average 18 person minutes were spent to find one millipede (range = 4 to 98 person minutes).

On two occasions Amplaria muiri were observed mating ( Fig. 27 ). On 15 July 2003, two individuals in the Whitewash Canyon section of Crystal Cave were observed mating on a fluffy sandy silt floor. On 13 October 2006, two individuals in the passage below the rope drop of Carmoe Crevice were observed mating on a gravel floor. A third individual was nearby and probably involved, as it was seen curled up so that its mouth was near its gonopods. Also on 13 October 2006, an unusual abundance of individuals were seen feeding on the seed of a California Bay Laurel, Umbellularia californica . Approximately thirty individuals were seen within 5 meters of the food item, and this abundance of prey also probably attracted a potential predator, the salamander Ensatina eschscholtzii ( Fig. 26 ).

Methods for habitat, abundance and life history: Temperatures were measured using an alcohol thermometer, and they were measured at locations in the same area of the cave the specimen was taken from, but not always exactly at that spot. Distances into the cave were calculated from survey stations measured during cave mapping, and these represent actual line of travel through the cave, not a straight line distance to the entrance that may be impossible to follow because it is through bedrock. The distance each organism was found into the cave was calculated for the search area the specimen was found in. The search area is bound by two survey stations, and if the closest survey station to the entrance is 10 meters into the cave and the farthest one is 20 m into the cave, the distance that specimen was found from the entrance is calculated at 15 m into the cave. The range that individual was found at is 10 to 20 m from the entrance. The distance between these two stations is an approximate measure of search area, in this case 10 linear meters of passage. In the cases of two long caves, Crystal Cave and Hurricane Crawl Cave, the distances were estimated based on personal experience in the cave. The search time was calculated as person minutes spent in that area, and this time also includes searching for non-millipede taxa, and time to collect specimens and record data. Furthermore, not every millipede seen was collected and these data are based on collected specimens, so the data may slightly overestimate rarity.

Notes: The caves of Sequoia National Park and Kings Canyon National Park, explored by JKK and her associates during a survey of cave fauna for the National Park Service, occur in five distinct clusters ( Fig. 23 ), which we have arbitrarily numbered 1–5 from north to south. Amplaria muiri has only been found in the caves of cluster 2. Unidentifiable striariids (males were not collected) were found in Lilburn Cave in cluster 1; these could also be A. muiri . As shown in Table 1 , millipeds attributable to A. muiri were collected in Bear Den Cave, Carmoe Crevice, Crystal Cave, Hurricane Crawl Cave, Lange Cave, Pet Cemetery Cave and Weisraum Cave. Males were not available from the last listed of these caves, but geography makes a good case that the specimens taken in Weisraum Cave are A. muiri . Specimens from Hurricane Crawl Cave and Weisraum Cave were moderately well-pigmented, but those from the other caves were pale yellowish white. Because no attempt was made to collect extensively in the forest litter and soil near the caves or elsewhere in the parks, it is not possible to say if A. muiri is cave-limited. The legs and antennae are not inordinately long, and while at the large end of the size spectrum of Amplaria species (except for the giant A. shastae ), a convincing picture of troglobiosis is not presented. The pale populations may represent permanent cave residents (troglophiles) but still may be in reproductive contact with surface populations.

Figures

FIGURE 23. Map of parts of Sequoia and Kings Canyon National Parks, Tulare Co., California, showing locations of caves studied in 2003 – 2004 by JKK and associates. Note that some of the cave names are shortened for ease of viewing on the map, typically the word “ Cave ” is left off. See text for the full names of caves in which Amplaria specimens were collected.

FIGURE 27. Amplaria muiri mating in Carmoe Crevice. Male above and slightly to the right; note use of encrassate anterior legs of male in clasping female.

FIGURES 4 – 6. Amplaria muiri, n. sp., male, scanning electron micrographs. 4. Coxae and trochanters of second legs of male, ventral view; p, seminal pore; cx, coxa; tr, trochanter. 5. Tibia and tarsus of first leg of male, showing modified setae on ventral surface, posterior view. 6. Modified setae from ventral surface of tibia.

FIGURES 7, 8. Amplaria muiri, n. sp., male. 7. Right gonopod, lateral view. 8. Right gonopod, mesal view.

FIGURES 9 – 14. Amplaria muiri, n. sp., male, scanning electron micrographs. 9. Gonopods, ventrolateral view; posterior to the right. 10. Gonopods, posterior view. 11. Gonopods, ventroposterior view. 12. Spine at corner of mentum and modified mandibular stipes, anterior view, ventral to the left. 13. Coxae of legpairs 3 – 5, anterior at the top. 14. Modified setae from coxa of leg 3.

FIGURES 15 – 18. New species of Amplaria, males, scanning electron micrographs. 15. A. muiri, ninth legpair, anterior view. 16 – 18. A. adamsi, n. sp. 16. Ninth legpair, anterior view. 17. Left gonopod, mesal view. 18. Right gonopod, lateral view.

FIGURES 24 – 26. 24. Amplaria muiri in Lange Cave. 25. Amplaria adamsii in Clough Cave. 26. Amplaria muiri ecology in Carmoe Crevice. To the right is an individual of the salamander Ensatina eschscholtzii, found near a large group of Amplaria muiri that were feeding on the seed of a California Bay Laurel (left). The salamander was not observed eating the millipeds, but likely preys on them.

Tables

TABLE 1. Collection localities for species of Amplaria in Sequoia and Kings Canyon National Parks. Cave names have been abbreviated; full names are given in the text.

Cave	Date	Collectors	Collection number	Species	Notes
Hidden	15-Nov-03	J. Krejca, V. Loftin	JKK-302-B1	Amplaria adamsi	male
Overhang	29-Apr-04	J. Krejca, P. Sprouse	JKK-22-C9	Amplaria adamsi	juvenile
Overhang	29-Apr-04	J. Krejca, P. Sprouse	JKK-25-C6	Amplaria adamsi	female
Overhang	29-Apr-04	J. Krejca, P. Sprouse	JKK-24-C4	Amplaria adamsi	male
Bear Den	17-Jul-03	J. Krejca, V. Loftin, S. Fryer	JKK-33-A3	Amplaria muiri	male
Bear Den	18-Jul-03	J. Krejca, V. Loftin, S. Fryer, C. Walck	JKK-333-A1	Amplaria muiri	2 males, female
Bear Den	1-May-04	J. Krejca, J. Snow, A. Snow, P Sprouse	JKK-43-C2	Amplaria muiri	2 penultimate instar males
Bear Den	1-May-04	J. Krejca, J. Snow, A. Snow, P Sprouse	JKK-40-C6	Amplaria muiri	5 males, 3 females
Carmoe Crevice	18-Nov-03	J. Krejca, V. Loftin	JKK-331-B2	Amplaria muiri	2 females, 4 juveniles
Carmoe Crevice	18-Nov-03	J. Krejca, V. Loftin	JKK-335-B1	Amplaria muiri	2 males, female, juvenile
Carmoe Crevice	18-Nov-03	J. Krejca, V. Loftin	JKK-330-B1	Amplaria muiri	3 males
Carmoe Crevice	5-Jul-04	J. Krejca, A. Gluesenkamp	JKK-1-D1	Amplaria muiri	2 males, female
Clough	19-Nov-03	J. Krejca, V. Loftin	JKK-341-B1	Amplaria adamsi	penultimate male
Clough	19-Nov-03	J. Krejca, V. Loftin	JKK-340-B9	Amplaria adamsi	2 males, 2 females
Clough	27-Apr-04	J. Krejca, P. Sprouse et al	JKK-3-C11	Amplaria adamsi	2 females
Clough	27-Apr-04	J. Krejca, P. Sprouse et al	JKK-2-C7	Amplaria adamsi	female
Clough	27-Apr-04	J. Krejca, P. Sprouse et al	JKK-1-C14	Amplaria adamsi	juvenile
Crystal	15-Jul-03	J. Krejca, V. Loftin, S. Fryer	JKK-003-A1	Amplaria muiri	15 specimens, type series
Crystal	15-Jul-03	J. Krejca, V. Loftin, S. Fryer, D. Boiano	JKK-005-A2	Amplaria muiri	male, female
Crystal	17-Jul-03	J. Krejca, V. Loftin	JKK-021-A1	Amplaria muiri	3 males, females
Crystal	17-Jul-03	J. Krejca, V. Loftin	JKK-027-A2	Amplaria muiri	5 females
Hurricane Crawl	16-Jul-03	J. Krejca, S. Fryer, A. Snow, V. Loftin	JKK-17-A5	Amplaria muiri	very small juvenile
Hurricane Crawl	16-Jul-03	J. Krejca, S. Fryer, A. Snow, V. Loftin	JKK-18-A3	Amplaria muiri	female
Hurricane Crawl	16-Jul-03	J. Krejca, S. Fryer, A. Snow, V. Loftin	JKK-10-A1	Amplaria muiri	female
Hurricane Crawl	9-Jul-04	J. Krejca, A. Gluesenkamp et al	JKK-47-D2	Amplaria muiri	2 males, female
Hurricane Crawl	9-Jul-04	J. Krejca, A. Gluesenkamp et al	JKK-47-D3	Amplaria muiri	very small juvenile
Hurricane Crawl	9-Jul-04	J. Krejca, A. Gluesenkamp et al	JKK-44-D2	Amplaria muiri	2 males, 2 females