A Re-Evaluation of the Millipede Genus Motyxia Chamberlin, with a Re

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Center for Systematic Entomology, Gainesville, Insecta Mundi Florida

September 1997

A re-evaluation of the millipede genus Motyxia Chamberlin, with a re-diagnosis of the tribe Xystocheirini and remarks on the bioluminescence (Polydesmida: Xystodesmidae)

Rowland M. Shelley North Carolina State Museum of Natural Sciences, Raleigh, North Carolina

Follow this and additional works at: https://digitalcommons.unl.edu/insectamundi

Part of the Entomology Commons

Shelley, Rowland M., "A re-evaluation of the millipede genus Motyxia Chamberlin, with a re-diagnosis of the tribe Xystocheirini and remarks on the bioluminescence (Polydesmida: Xystodesmidae)" (1997). Insecta Mundi. 280. https://digitalcommons.unl.edu/insectamundi/280

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 331

A re-evaluation of the millliped genus Motyxia Ghamberlin, with a re-diagnosis of the tribe Xystcseheirini and remarks on $he bislumi- nescenee (Polydesmida: Xystsdesmidae)

Rowland M. Shelley North Carolina State Museum of Natural Sciences, P. 0. Box 29555, Raleigh, North Carolina 27626-0555

Abstract: Motyxia Chamberlin is comprised of eight species of bioluminescent xystocheirine millipeds in which the gonopodal solenomere arises at different positions, from basally and subbasally on the acropodite to being fused with the companion acropodal branch and detachingproximad or near midlength. Previous synonymies ofilrnplocheir Chamberlin and Lurrril~odesrnl~sLoomis and Davenport under Motyria are confirmed as is its assignment to the tribe Xystocheirini, which is redefined. Component species are Ak Izerr~aChamberlin, the type species, r~rortica Chamberlin, sequoiae (Loomis and Davenport), tularea (Chamberlin), sequoia (Chamberlin), pi01 Chamberlin, pol.l.ecta Causey and Tiemann, and tierr~ariu~i Causey. Motyxia sequoia is comprised of two races, the nominate and sequoia nlia Causey andTiemann; sequoia ollae Causey andTiemannis properly a subspecies of tularea. A4otyxiapio1,formsecca is an invalid name without standingin nomenclature, and M. tejorla Chamberlin, andM. erpalrsa and exilis, both by Loomis, are placed in synonymy under M. IILOII,~~~,the oldest name for the southernmost species, as Polydesrilus dissectzrs Wood is referrable to Xystocheir Cook. The bioluminescence is a continuous, neon-white glow of the entire dorsal surface including the antennae and 1egs.Its visibility at night suggests a warning function analogous to aposematic coloration. The phenomenon may observe a circadian rhythm, and controlled photoperiod experimentation may be productive.

both subspecies of M. tularea; it is fused basally to Introduction the companion branch and detaches proximad in M. sequoia alia Causey and Tiemann; and it is The dominant xystodesmid milliped genus in fused and detaches near midlength in M. s. sequoia the southern Sierra Nevada, south of Sequoia Na- (Chamberlin), the type species ofAlnplocheir Cham- tional Park, is Motyxia Chamberlin. It ranges west- berlin, M. pior Chamberlin, and M. porrecta. The ward to the Pacific Ocean through the Tehachapi intermediate positions between the extremes in the and Santa Monica Mountains and is the southern- type species of Motyxia and Ainplocheir confirm most representative of the family in California and Causey and Tiemann's assignment (1969) of the western North America (Figs. 1, 2). The only genus latter to synonymy, as Motyxia is the oldest genus- of bioluminescent millipeds, Motyxia, was revised group name for this assemblage. The third nominal by Causey and Tiemann (1969), but the cyphopods genus, Lu~~zit~odes~nusLoomis and Davenport, is were not characterized, and two nomenclatural an obvious synonym because the gonopods of its changes are necessary. Motyxia sequoia ollae Cau- type species conform to the pattern in M. kerna. sey and Tiemann is actually a race of M. tularea Motyxia encompasses relatively large-bodied xys- (Chamberlin), and M. lnoltica Chamberlin, the old- todesmids that are distinguished somatically by est of four synonyms, is the correct name for the their orange base color, absence of dorsal papilla- southernmost species, as Polydeslnus dissectus tion, and orientation of the anterior paranota, Wood is referrable to Xystocheir Cook (Shelley which angle anteriolaterad on segments 2-5 (Figs. 1996). 3-4). Gonopodal similarities between M. s. sequoia Motyxia comprises a spectrum of forms in which and Xystocheir reducta (Causey) indicate generic the origin of the solenomere undergoes a complete affinity and contribal status, so the Xystocheirini transition regarding its position relative to the must be rediagnosed to accommodate Motyxia, companion acropodal branch. The solenomere orig- whose species lack the lobes on the 3rd coxae of inates basally on the acropodite and is widely males and the 2nd of females that are present on segregated from the companion branch in M. Kertta other tribal components (Hoffman 1980). Because Chamberlin, the type species, M. moltica, and M. of apparent homologies, I assign the companion sequoiae (Loomis and Davenport); it arises basally acropodal projection in Motyxia, lying caudadllat- and is narrowly segregated from the companion erad to the solenomere, the same label as in Xys- branch in M. tiemanni Causey; it arises subbasally tocheir, "process 'B"' (Shelley 1996). and is moderately segregated from the latter in 332 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Fig. 1. Distribution ofMotyxia. A smooth curve is drawn around range extremes in all directions, and the question mark denotes the dubious record ofM. ~no~~icafrom Riverside County. Fig. 2. Distributions ofspecies ofMotyxia. asterisks, M. rnol~ica;diamonds, M. kerlaa; solid stars, M. sequoiae; solid squares, M. t. tularea; open squares, M. ttula,i.eaollae; triangles, M. s. sequoia; inverted triangles, M. sequoia alia; X's, M. sequoia intergrades; dots, M. pior (thecircle indicates the sight record from Crystal Cave);half-shaded dots, M. porrecta; open stars, M. tie~nalrlai. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 333

Figs. 3-8. Motyxza I~ZOILZCU.3, head and tergites 1-4of a male from Woodford, dorsal view. 4, tergites 9-11 of the same, dorsal view. 5, left gonopod of the same, medial view. 6,the same, lateral view 7, acropodite of the same, caudal view. 8, left cyphopodof female from the same locality, caudal view. B, process "B; CV, caudal valve; 0, operculum; PFP, prefemoral process; S, solenomere. Scale lines for figs. 3-4 = 1.00 mm; line for other figs. = 1.00 mm for each. 334 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

The dominant North American polydesmoid personal observations, and suggesting a possible family, occurring in three areas of the continent avenue for productive research. Acronyms of sourc- (Shelley 1987) - east of the Central Plains, along es of preserved study material are as follows: the Paclfic Coast from Los Angeles to southern Alaska and inland to western Montana, and from AMNH - American Museum of Natural History, New southern Texas and New Mexico to El Salvador - York, New York. the Xystodesmidae has been the subject of numer- CAS - California Academy of Sciences, San Francisco. ous systematic studies since 1940. This contribu- FSCA - Florida State Collection of Arthropods, Gaines- ville. tion completes modern treatments of the west- LACMNH - Los Angeles County Museum of Natural Nearctic fauna that began with the revision of History, Los Angeles, California. Orophe Chamberlin and the proposal of the sub- NCSM - North Carolina State Museum of Natural Sci- family Orophinae (Hoffman 1964). Buckett and ences, Raleigh. Gardner (1968a, b) revised Harpaphe Cook (Harpa- NMNH - National Museum of Natural History, Smith- phini) and Wa~nohiaChamberlin (Xystocheirini), sonian Institution, Washington, D. C. and subsequently proposed the new genera UCB - Essig Entomological Museum, University of Cal- Anolnbrocheir (Xystocheirini) (1969a, b) and ifornia at Berkeley. UCD - Bohart Entolnological Museum, University of (Chonaphini) (1969~).After Causey Metaxycheir California at Davis. and Tiemann's (1969) study of Motyxia, eleven UCR - Entomology Department, University of Califor- years elapsed before Hoffman (1980) established nia at Riverside. the Harpaphini and Xystocheirini, and reduced the Orophinae and Chonaphinae Verhoeff (1941) to Literature Review tribal status in the subfamily Xystodesminae. Thir- teen years later, I (Shelley 1993a, b, c) updated The history of Motyxia began with the paper by Orophe, proposed Thrinaphe (Harpaphini), and Chamberlin (1941), in which he described Xys- revised Isaphe Cook (= Hybaphe Coolr) (Harpaphi- tocheir sequoia, from Sequoia National Park, and ni). These were followed by revisions of the tribes erected Motyxia. for M. her~za,from Kern, and M. Chonaphini and Sigmocheirini (Shelley 1994,1995a) pior, from Tulare, counties. Chamberlin (1944,1947) and the proposal of Parcipro~nus(Xystocheirini) proposed M. ~nonica,from the Santa Monica Moun- (Shelley 199513). The companion p~tblicationon tains, and M. tejo~za,from Fort Tejon, respectively; Xystocheir (Shelley 1996) addresses the last unre- he (Chamberlin 1949) proposed X. tularea for vised "western" genus, leaving only two unstudied another species from Tulare County and estab- genera in the United States -- the "eastern" taxa lished Alnploclzeir for X. sequoia. Loomis and Dav- Apheloria. and Nannaria, both authored by enport (1951) proposed Luminodesmus for L. se- Chamberlin. Shelley (1989) described Rhysodes~nus quoiae, also from Tulare County, and Davenport et clzisosi, from Big Bend National Park, Texas, and al. (1952) studied its ecology, life history, and (1987, 1992) addressed Ste~zodes~rsustuobitus bioluminescence, one of the few works on the "biol- (Chamberlin), in southern Texas and New Mexico, ogy" of any xystodesmid. Loomis (1953) described these being representatives of the "Meso-American M. exilis, from the Tehachapi Mountains, and M. fauna." This reexamination of Motyxia supple- expama, from Fort Tejon, the second nominal spe- ments Causey and Tiemann's revision (1969) by cies to be proposed from this site. Chamberlin and providing a historical section, an improved key to Hoffman (1958) incorporated all of the preceding species and subspecies, cyphopod illustrations, and taxa into the North American Checklist and placed standardized gonopod drawings in medial and lat- Lulni~zodes~nusin synonymy under Xystocheir. eral views that are more readily compared and In recent works, Causey (1960) described M. interpreted than their SEM photos. Species ac- tielnal~~~i,from Kern County, but Buckett (1964) counts include synonymies, listings of type speci- omitted this species from his list of California mens, diagnoses, statements on variation and millipeds. Causey and Tiemann (1969) revised distribution, and new localities (the initials DLT Motyxia, placed A~nplocheirand Lu~ninodes~nusin identify samples collected by Darwin L. Tiemann). synonymy, and transferred X. dissecta and tularea, I also supplement previous reports on the A. sequoia, and L. sequoiae into the genus. They bioluminescence (Loomis and Davenport 1951; also placed M. ~no~zica,tejo~za, expama, and exilis Davenport et al., 1952; Causey and Tiemann 1969, in synonymy under dissecta; proposed M. porrecta 1970) by summarizing current knowledge, adding for another species from Kern County; and divided INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997

M. sequoia into three races, the nominate and the Components. Xystocheir Cook, Warnokia new subspecies, M. s. alia and s. ollae. The same Chamberlin, Motyxia Chamberlin, Artombrocheir authors (Causey and Tiemann 1970) summarized Buckett and Gardner, Parciprornu Shelley. the distribution, biology, and bioluminescence of the species of Motyxia, and Jeekel (1971) included Genus Motyxia Chamberlin Motyxia, Alnplocheir, and Lurni~todesinusin the Motyxia Chamberlin, 1941:15. Chamberlin and Hoff- Nomenclator. In the final relevant work, Hoffman man, 1958:38. Buckett, 1964:9. Causey and Tie- mann, 1969:14-20. Jeekel, 1971:274. Hoffman, (1980) assigned Motyxia, with the synonym 1980: 157. Lurnirzodesmus, to the new tribe Xystocheirini and A~~zplocheirChamberlin, 1949:97. Chamberlin and Hoff- revived Arnplocheir as a monotypic genus for X. man, 1958:17. Buckett, 1964:7. Jeekel, 1971:247. sequoia, the type species. Hoffman, 1980:157. Lt~~ni~zodes~~zusLoomis and Davenport, 1951:270-271. Tribe Xystocheirini Hoffman Jeekel, 1971:271.

Diagnosis. Moderate to large size Xystodesmi- Type species. Of Motyxia, M. lzerrza Cham- nae with smooth and glossy to variably papillate berlin, 1941, by original designation; ofAlnplocheir, dorsums, pustulate caudad in some species; sides of Xystocheir sequoia Chamberlin, 1941, by original collum gently curved or folded and uplifted at designation; of Lu~nir~odesmus,L. sequoiae Loomis midlength; caudolateral paranotal corners either and Davenport, 1951, by original designation. broadly rounded and not prolonged or subacumi- Diagnosis. Moderate-size to large, biolumi- nate and extending slightly or strongly caudad, nescent Xystocheirini with orange base color; occasionally curving mediad and appearing unci- metatergites smooth and glossy, without papilla- nate; 3rd male coxae with or without subconical, tion; sides of collum not uplifted; caudolateral para- anteriorly directed lobes; 2nd female coxae with or notal corners broadly rounded, not prolonged without long, conical lobes overlying cyphopodal caudad; 3rd male and 2nd female coxae without aperture; gonopodal aperture generally ovoid, cau- lobes; gonopodal prefemur generally upright, usu- dal margin with at most only slight caudal exten- ally sparsely hirsute, with or without distolateral sion; gonosternum strong, oriented along longitu- lobe; prefemoral process variable but usually long dinal, anteriorlposterior body axis; coxa flattened, and broad, upright, sinuate, or curving anteriad or without apophysis, usually with one or two mac- caudad, entire or divided into variably broad medi- rosetae, occasionally with field; prefemur generally al, and narrower lateral, subbranches, former with sublinear and massive, densely or sparsely hirsute, or without basal spur; solenomere variably long, with or without distolateral lobe extending beyond arising either basally from acropodite as distinct, acropodal articulation; prefemoral process usually acicular projection, curving strongly or gently strong and clearly detached from stem, occasional- caudolaterad, widely or narrowly segregated from ly vestigial or absent, located either on anterio- process "B,or fused basally to latter and detaching lateral corner, on medial surface short of latter, or proximad or near midlength; process "Bvarying in caudolateral corner; acropodite highly variable and length, breadth, and configuration, broad and clearly demarcated from prefemur, consisting of sublaminate or narrow and subacicular, upright, either an undivided/shallowly divided structure, sinuate, or curving variably anteromediad; cypho- either upright, curving in sigmoid configuration, or pods oriented transversely in aperture, valves large narrow proximad and expanded distad, or from two and moderately hirsute, unequal with anterior to four terminal projections of varying configura- valve larger, receptacle varying in size, hirsute or tions; solenomere either a separate branch arising glabrous, absent from one species, operculum small from base to midlength of acropodite or a shorter and moderately hirsute. projection arising proximad to near midlength of Distribution. Occupying two segregated ar- caudal acropodal branch (process '3");cyphopods eas in southern California, a broad one extending in usually oriented transversely in aperture, valves a generally northlsouth direction from the western usually densely hirsute, often with bent hairs, with side of Sequoia National Park, in northern Tulare hirsute operculum and hirsute or glabrous recepta- County, through the southern periphery of the cle located on dorsolateral and dorsomedial corners Sierra Nevada and the Tehachapi Mountains to of valves, respectively, with or without moderately Fort Tejon, in southern Kern County, and a small, sclerotized, glabrous, lateral accessory body. disjunct area near the Pacific Coast in the Santa 336 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Monica Mountains, Los Angeles County (Fig. 1). gonopod prefemora are sparsely hirsute with only Species distributions are mutually exclusive except scattered hairs on the stem and a light cluster or no for overlap between M. kerl~a,tularea, and sequoi- hairs basally around the pit of the prostatic groove. ae in central and southern Tulare County (Fig. 2). The cyphopod receptacle is absent from this spe- The species with undivided prefemoral processes cies, glabrous in M. tularea, porrecta, and tielna7~- (three total telopodal projections) occur on the IL~,and hirsute in the remaining species; however, peripheries -- M. pior on the north, and M. tielna7~- the setation is notably sparse in M. Izerl~a,sequoiae, IL~,porrecta, and 7no7~icaon the south -- while those andpior. The glabrous receptacles distinguish these with divided processes (four total branches) occur species from all other western xystodesmids, and centrally. This pattern implies that loss of the M. 1no7~icais the only Pacific representative lack- second prefemoral subbranch has occurred twice ing the structure. and that the three-branched condition in M. pior is Many specimens of Motyxia were collected in convergent with those in the other species. the 1960's by Darwin L. Tiemann (Shelley 1995a, Species. Eight, two divided into two geograph- 1996); some were coated with shellac or another ic races each. The occupied area in California has substance and placed on insect pins at the LACM- been thoroughly sampled (Tiemann 1963, 1964), NH, and most of the others were deposited in Dr. and all species have probably been discovered. Causey's private collection, which was transferred However, future sampling may disclose intercon- to the FSCA after her death in 1979. The pinned necting forms among the species with divided specimens are unusable except for external, i7~situ prefemoral processes. Motyxia sequoia is definitely examinations and measurements, and I present composed of two races, as the nominate subspecies tabulated data for all taxa in the variation sections, and M. s. alia are linked by anatomically and with localities listed in general north to south geographically intermediate forms in the vicinity of sequences. Clough Cave in the southwestern corner of Sequoia National Park. I also recognize two races of M. Key to species and subspecies of Motyxia tularea because of their narrow separation and the close similarity of their gonopods; while none are 1. Prefemoral Drocess divided into medial and lateral subbranches, telopodite with 4 terminal projec- available, intergrades are presumed to occur in the tions (Figs. 9-11, 13-15, 17-19, 21-26, 28-31) .. lacuna. Motyxia Izer7~aand M. sequoiae are also ...... 2 very close anatomically, the chief difference being - Prefemoral process not divided, telopodite with 3 the apical configuration of the medial prefemoral terminal projections (Figs. 5-7, 32-36-38, 40-41) subbranch; however, the intervening area has been ...... 7 reasonably well sampled without the discovery of intergrades, so I interpret this difference as evi- 2. Medial subbranch of prefemoral process with variable dence of reproductive isolation. Beyond these ex- basal spur; solenomere fused basally to process amples, I think that meticulous sampling in "B" (Figs. 25-26,28-31) sequoia (Chamber1in)S - Medial subbranch of prefemoral process without uncollected areas in central Tulare County may basal spur; solenomere arising basally or sub- reveal anatomically intermediate forms that join basally from acropodite (Figs. 9-11, 13-15, 17- all the four-branched species, because their gono- 19, 21-24) ...... 4 podal differences are so small asto be easily bridged. For example, M. sequoia is diagnosed primarily by 3. Detached part of solenomere relatively long, subequal the basal spur on the medial prefemoral subbranch, in length to distal part of process "B"; latter and but this structure varies in size, and a vestigial medial subbranch of prefemoral process rela- spur, coupled with a more proximal detachment of tively narrow for entire lengths, sides tapering the solenomere from process "B", would blur the smoothly and continuously to subacuminate tips (Figs. 25-26); vicinity of Hammond, Tulare distinction between M. s. alia and M. tularea ollae, County ...... s. sequoia (Chamberlin) and imply that M. sequoia and tularea are conspe- - Detached part of solenomere relatively short, less cific. Thus, while the present material indicates than half as long as distal part of process "B"; eight species, the actual number may be only four latter and medial subbranch of prefemoral pro- or five. cess relatively broad to subspatulate for at least Remarks. Aside from bioluminescence, Motyx- part of length, sides expanding either proximad ia is characterized by a loss of hairs on both the or distad, apically broad or rounded (Figs. 28- gonopods and cyphopods. Except for M. mo7~ica,the 29); Three Rivers to Balch Mountain Park, INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 337

Tulare County ...... pital Rock, Sequoia National Park, Tulare ...... s. alia Causey and Tiemann County ...... pior Chamberlin - Process "B" subspatulate, extending ventrad in sinu- 4. Solenomere arising basally on acropodite, curving ate configuration; detached part ofsolenomere strongly caudolaterad, overlapping margin of short but distinct, clearly projecting from mar- process "B"; latter extending caudolaterad ba- gin of process "B(Figs. 36-39);Kern River Val- sally then curving anteromediad and extending ley, Kern County ...... ventrad (Figs. 11-15) ...... 5 ...... porrecta Causey and Tiemann - Solenomere arising subbasally on acropodite, either upright and extending directly ventrad or curv- Motyxia monica Chamberlin ing only apically and at most only slightly Figs. 3-8 caudolaterad; process "B" generally upright (Figs. 17-19, 21-24) tularea (Chamberlin) ..... 6 Motyxia rno7zica Chamberlin, 1944:57, figs. 1-3. Cham- berlin and Hoffman, 1958:39. Buckett, 1964:9. 5. Medial branch of prefemoral process apically expand- Motyxia tejo~zaChamberlin, 1947:25, figs. 4, 4a. Cham- ed and upright (Figs. 9-10); Cold Springs Sad- berlin and Hoffman, 1958:39. Buckett, 1964:9. New dle and White River vicinity, Tulare County, to synonymy. northern fringe of Kern County ...... Motyxia expalzsa Loomis, 1953:422, fig. 19. Chamberlin ...... Izer7za Chamberlin and Hoffman, 1958:39. Buckett, 1964:9. New syn- - Medial branch of prefemoral process apically sig- onymy. moid, curving caudolaterad (Figs. 13-14); Camp Motyxia exilis Loomis, 1953:422, fig. 20. Chamberlin Wishon to Coy Flat, Tulare County ...... and Hoffman, 1958:39. Buckett, 1964:9. New syn- ...... seq~soiae(Loomis and Davenport) onymy. Motyxia dissecta (Nec Wood) Causey and Tiemann, 6. Medial branch of prefemoral process with caudal 1969:21-22, figs. 2-3, 8, 10-12. margin indented apically, tip acuminate (Figs. 17-19); Quaker Meadows, Tulare County, to Type specimen. Male holotype (NMNH) col- northern fringe of Kern County ...... lected by R C. Stebbins, 4 March 1944, at "Madelia ...... (Chamberlin) tularea tularea Canyon," Sherman Oaks, Santa Monica Moun- -Medial branch of prefemoral process apically expanded and rounded (Figs. 21-22, 24); Camp Wishon tains, Los Angeles County. The vial label and orig- to Deer Creek Road south of Tule River Indian inal description state, "Meadow Canyon," which is Reservation ...... believed to be a lapsus for Madelia Canyon. Made- ...... tztlarea ollae Causey and Tiemann lia Avenue is a short street off Valley Vista Boule- vard west of Beverly Glen Boulevard on the north 7. Solenomere arising basally on acropodite (Figs. 5, 7, slope of the Santa Monica Mountains in Sherman 40) ...... 8 Oaks, between Mulholland Drive and the Ventura - Solenomere fused basally to process "B, detaching Freeway. proximad to near midlength (Figs. 33-34,37-38) Diagnosis. Prefemoral process entire, broad basally, narrowing smoothly and continuously to 8. Solenomere widely segregated basally from process narrowly rounded tip, curving broadly laterad; "B"; latter extending sublaterad basally then solenomere arising basally from acropodite, curv- curving broadly anteromediad and extending ing broadly caudolaterad, directed toward midlength ventrad; cyphopod without receptacle (Figs. 7- of, but not overlapping, process "B; latter widely 8); vicinity of Breckenridge, Kern County, to segregated basally from solenomere, extending Santa Monica Mountains, Los Angeles County sublaterad basally then curving broadly antero- ...... rno~zicaChamberlin mediad and extending curvilinearly ventrad, sides - Solenomere narrowly segregated basally from pro- expanding basally then narrowing smoothly and cess "B"; latter upright, extending directly ventrad; cyphopod with receptacle (Figs. 40, 42); continuously to subacuminate tip; cyphopod with- Greenhorn Mountains from Tulare County Line out receptacle (Figs. 5-8). to Evans Flat, Kern County ...... Variation. The gonopods of M. monica are ...... tierna7a7zi Causey quite uniform, the only observable variation being slightly less or greater curvatures of the projections 9. Process "B broad and semilunar; detached part of thanillustratedinfigs. 5-7. Measurements of pinned solenomere minute and blunt, barely projecting adults are summarized in table 1; sexes are sub- beyond margin of process "B (Figs. 32-24); equal in size, and on the average, individuals be- Woodlake Valley east to Crystal Cave and Hos- 338 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI come shorter and narrower to the south in the range. Type specimen. Male holotype (NMNH) collected Distribution. Occurring south of the Kern by S. and D. Mulaik, 19 March 1942,7 mi (11.2 km) River in two apparently disjunct areas of unequal N Glenville, Kern County. size, the larger extending from Breckenridge Moun- Diagnosis. Prefemoral process divided basally tain through the Tehachapi Mountains to Fort into narrow, linear lateral, and broad, expanded Tejon, Kern County, and the smaller being near the medial subbranches, former 314 as long as latter, Pacific Ocean, some 50 mi (80 km) to the south, in not bowed anteriad, latter entirely upright, apical- the Santa Monica Mountains, Los Angeles County. ly expanded and broadly rounded; solenomere aris- Known elevations range from around 700 to 7,000 ing basally from acropodite, curving strongly ft. The intervening area of eastern Ventura Coun- caudolaterad, overlapping and extending slightly ty, in the Los Padres National Forest around Pyr- beyond margin of process "B; latter widely seg- amid and Pinu Lakes, is plausible for M. inonica. regated basally from solenomere, extending direct- A new locality is available from the Santa ly laterad basally then curving strongly anterome- Monica Mountains, but metropolitan Los Angeles diad and extending sublinearly ventrad, sides ex- now covers much of this range, and south-facing panding slightly distal to curve then narrowing to canyons like Beverley Glen are virtually complete- narrowly rounded tip; cyphopod with hirsute re- ly urbanized. I drove the length of Beverley Glen ceptacle (Figs. 9- 12). Boulevard in 1989, but with continuous houses and Variation. The apical expansion of the medial private property on both sides, could not find any- prefemoral subbranch is greater in scattered males, place to search. Western facing Topanga Canyon, and the solenomere curls dorsad apically in a few in the Santa Monica Mountains National Recre- males from the vicinity of White River. Measure- ation Area, is a more plausible site, but I could not ments of pinned adults are summarized in table 2; find any specimens in three visits. Searches to the the few available females are larger in both dimen- east, in the San Gabriel Mountains of Los Angeles sions than average males from the same sites, but and San Bernardino counties, have not revealed no geographic trends are evident. the milliped, but a dubious sample is available from Distribution. Southern Tulare County from Riverside, Riverside County, some 80 mi (128 km) the vicinity of Cold Springs Saddle to the adjacent east southeast of the Santa Monicas. This city is northern fringe of Kern County, extending west- inconsistent with the known, coherent ranges of ward to the vicinity of White River along Arrastre both the genus and species; specimens have not Creek Road, distances of around 10.8 mi (17.3 km), been collected there again; and the precise location northlsouth, and 12.8 mi (20.5 km), eastlwest. in Riverside is unknown. The desert habitat at Specimens were examined from the following new Riverside is wrong for M. inonica, whose known localities: Tulare Co., 7.3 mi (11.7 km) NE White environments are more moist with an abundance of River, 68M, 15F, 21 February 1965, DLT (FSCA, trees, so the locality almost surely is wrong. Spec- LACMNH); 1 mi (1.6 km) "below" Sugarloaf Mtn. imens were examined from the following new local- Park, 18M, 8F, DLT (FSCA); and 7 mi (11.2 km) ities: Los Angeles Co., Santa Monica Mts., Topan- Greenhorn Mtn. Peak, along CA hwy. 15, M, F, 1 ga Cyn. at Greenleaf Cyn., 10M, 5F, 8 December June 1984, T. Schackman (NCSM). 1966, DLT (LACMNH), and Beverly Glen Cyn., 2M, F, March 1953, collector unknown (NMNH). Motyxia sequoiae (Loomis and Davenport) Dubious Record. Riverside Co., Riverside, Figs. 13-16 3M, Fall 1953, L. D. Anderson (UCR). Lunzii~odesrnrssseqz~oiae Loomis and Davenport, Remarks. Shelley (1996) showed that Wood's 1951:271-272, fig. 1. name, dissecta, is correctly referrable to Xystocheir Xystocheir sequoiae: Chamberlin and Hoffman, 1958:54- Cook and that the purported locality, Fort Tejon, is 55. Buckett, 1964:lO. wrong. Consequently, M. inonica, the oldest of the Motyxia seq~~oiaeCausey and Tiemann, 1969:31, figs. four available species-group names, is the correct 17-18. name for this species. Type specimens. Male holotype and female Motyxia kerna Chamberlin paratype (NMNH) collected by D. L. Tiemann, 12- Figs. 9-12 14 May 1950, above Camp Nelson, Sequoia Nation- Motyxia kerna Chamberlin, 1941:15, pl. 3, fig. 29. Cham- al Forest, Tulare County. berlin and Hoffman, 1958:39. Buckett, 1964:9. Cau- sey and Tiemann, 1969:25, figs. 4, 19-20. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 339

rowing to narrowly rounded tip; cyphopod with hirsute receptacle (Figs. 13-16). Variation. The lateral prefemoral subbranch is bowed strongly anteriad in males from Camp Wishon but moderately to lightly so in those from Camp Nelson and Belknap Campground. Measure- ments of pinned adults are summarized in table 3; on the average, females are larger in both dimensions than males from the same site, but no geographic trends are evident. Distribution. Known only from a small, sym- Figs. 9-12. Molyria kerns. 9, left gonopod ofa Inale from 3.3 metrical area, 4.4 mi (7.0 Irm) in both dimensions, mi (5.3 km) SE White River, medial view. 10, the same, lateral in the headwaters of the Middle Fork of the Tule view. 11, acropodite of the same, caudalview. 12, left cyphopod of River. It extends, northlsouth, from Camp Wishon afemale from the samelocality, caudalview. L, lateralsubbranch to coy~l~t and, east/west, from cedar slopeto the ofprefemoral process; M, medial subbranch ofprefemoral process; R, receptacle; other abbreviations as in figs. 3-8. Scale line = 1.00 crossing of Moorehouse Creek by highway 190. No mln for all figs. new localities are known.

Diagnosis. Prefemoral process divided basal- Motyxia tularea (Chamberlin) ly into relatively narrow, linear lateral and moderately broad, expanded medial subbranches, former Diagnosis. Prefemoral process divided basal- 314 as long as latter, not bowed anteriad, latter ly into narrow, linear lateral and broad, expanded, generally upright but curving caudolaterad distal- subspatulate medial subbranches, former 112-314 ly in sigmoid configuration, sides narrowing prox- as long as latter, bowed slightly anteriad, latter imal to curve, expanding greatly distad then nar- upright, apically broad or narrow; solenomere aris- rowing to blunt tip; solenomere arising basally ing subbasally from acropodite, generally upright, from acropodite, curving strongly caudolaterad, at most curving only slightly caudolaterad, barely overlapping but not extending beyond process "B"; overlapping process "B";latter generally upright, latter widely segregated basally from solenomere, sides expanding proximad, narrowing distad to angling sublaterad basally then curving broadly narrowly rounded tip; cyphopod with glabrous re- anteromediad and extending curvilinearly ventrad, ceptacle. sides expanding strongly distal to curve then nar- Remarks. Intergrades are not available, but two closely similar and narrowly segregated forms, 340 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Figs. 13-16. Motyxio sequoioe. 13, left gonopod of a inale from Belknap Campground, medial view. 14, the same, lateral view. 15, acropodite of the same, caudal view. 16, left cyphopod of a female from the same locality, caudal view. Abbreviations as in figs. 3-8,9-12. Scale line = 1.00 lnln for all figs. differing primarily in the apical configurations of date in 1930 on Sugarloaf Mountain, Tulare Coun- the medial prefemoral subbranch and process "B", ty. are interpreted as representing races of this spe- Diagnosis. Medial subbranch of prefemoral cies. process apically narrow and acuminate, indented on caudal margin and curving caudad; process "B Motyxiatuzarea tuzarea (Chamberlin)y new entirely upright, sides narrowing gradually distal status to midlength (Figs. 17-20). Figs. 17-20 Variation. The medial prefemoral subbranch is curved more strongly caudad and has two tiny Xystocheir tz~lareaChamberlin, 1949:101, fig. 27. Cham- berlin and Hoffman, 1958:55. Buckett, 1964:ll. apical teeth in males from Tyler and Mead- Motyria tularea Causey and Tiemann, 1969:32, figs. 25- OWS; those from Capinero Creek have the 26 teeth but are curved as in the drawings. Measure- ments of pinned adults are summarized in table 4; Type specimens. Male holotype, female allo- On the average, females are larger than males from type, and two male and one female paratypes the same sites, and there is a general trend toward (NMNH) collected by E(. Denner on an unknown shorter and narrower individuals to the south in the range. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 341

Figs. 17-20. Motyxia tularea tularea. 17, left gonopod of a male from Long Meadow, medial view. 18, the same, lateral view. 19, acropodite of the same, subcaudal view. 20, left cyphopod of a female from the same locality, caudal view. Abbreviations as in figs. 3-8, 9-12. Scale line = 1.00 mm for all figs.

Motyxia tularea ollae Causey and Tiemann, new combination Figs. 21-24

Motyxia sequoia ollae Causey and Tiemann, 1969:29-30, Distribution. A sublinear area extending from figs. 6, 27-28. Quaker Meadows in Tulare County to the adjacent northern fringe of Kern County in the Greenhorn Type specimens. Male holotype, female allo- Mountains, a distance of some 21.3 mi (34.1 km). type, and 15 male and 9 female paratypes (FSCA) The race occurs sympatrically with M. kerns in the collected by D. L. Tiemann, 28 February 1965, 5.4 Sugarloaf Mountain area in southern Tulare Coun- mi (8.6 km) W of the Tule River Indian Reservation, ty. Specimens were examined from the following Tulare County. For data on other paratype Sam- new localities: Tulare Co., Long Mdw. W of ples>see cause^ and Tiemann (1969). Johnsondale, 19M, 2F, 28 May and 16 June 1961, Diagnosis. Medial subbranch of prefemoral DLT (FSCA, MCMNH); Tyler ~d~.,5~,6~, 10 process apically broad and expanded; process "B" june1962, DLT (MCMNH); 0.9 mi (1.4 km) N curving slightly caudad, sides narrowing rapidly Kern Co. Line, Greenhorn Mts., 30M, 4F, 15 May distal to midlength (Figs- 21-24). 1965, DLT, L. W. Nichols (LACMNH); and Kern/ Variation. The medial prefemoral subbranch Tulare Co. Line, 5M, F, 15 May 1965, DLT, L. W. curves more strongly caudad in males from below Nichols (LACMNH). Kern Co., 0.2 mi (0.3 km) S Camp Wishon, Deer Creek Road at pothole creek, Tulare co. Line, 28~,4F, DLT, L. W. ~i~h~l~and 4.1 mi (6.6 km) east of the entrance to the Tule (LACMNH). River Indian Reservation (Fig. 24). These individ- 342 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI uals vaguely resemble M. kerns but are distin- nate; solenomere fused basally to process "B", aris- guished by the upright solenomere. Measurements ing proximad to near midlength of latter, detached of pinned adults are summarized in table 5; except part varying in length and breadth, subequal to, or for the one from Pothole Creek, females are larger much shorter than, distal part of process "B",ex- in both dimensions than males from the same sites, tending subparallel to latter or diverging slightly, and there is a general trend toward larger individ- apically acuminate; distal part of process "B"long uals to the south in the range. and variably broad, narrow to subspatulate, gener- Distribution. From just below Camp Wishon, ally upright, either reflexed caudad apically or on the North Fork of the Middle Fork of the Tule curving or leaning gently to broadly anteriad, api- River, southward to Deer Creek Road at Pothole cally broad or acuminate; cyphopod with hirsute Creek, south of the Tule River Indian Reservation, receptacle. westward to 5.4 mi (8.6 km) west of the entrance to Remarks. Motyxia sequoia is comprised of the latter. Dimensions are approximately 16.8 mi peripheral races connected by geographically and (26.9 km), northlsouth, and 11.1 mi (17.8 km), east1 anatomically intermediate forms. Chamberlin's west. Specimens were examined from the following original assignment (1941) to Xystocheir was reason- new localities: Tulare Go., along Dunn R. Fire Tr., able, because its two-branched acropodite demon- 2.5 mi (4.0 km) "below" Camp Wishon, 12M, 28 strates the pattern of this genus, particularly that December 1963, DLT (FSCA); and entrance to Tule of X. reducta (Causey), and the complex prefemoral R. Indian Res., 4M, 12F, 28 February 1965, DLT process is reminiscent of that in X. stolol~ifera (FS CA) . Shelley (Shelley 1996). However, without dorsal Remarks. After restudying all of Causey and papillation and lobes on the 3rd male and 2nd Tiemann's material, now at the FSCA, I agree with female coxae, sequoia was misplaced in Xystocheir, their conclusion as to the existence of the species- which may be why Chamberlin (1949) established group taxon that they named ollae. However, it is Arnplocheir to accommodate it. As explained previ- a race of M. tularea instead of M. sequoia, both of ously, Alnplocheir is a synonym of Motyxia, and which possess two prefemoral and two acropodal because of the gonopodal similarity between M. projections. The lateral prefemoral subbranch is sequoia and X. reducta, Motyxia is clearly contribal not bowed anteriad as strongly as in M. sequoia, with Xystocheir. and the medial one lacks the basal spur, the key diagnostic feature of the latter. Likewise, the distal Motyxia sequoia sequoia (Chamberlin) configurations of the medial prefemoral subbranch Figs. 25-27 and p-rocess "B" conform to the general shape of the structures in M. tularea and differ from those in Xystocheir seqz~oiaChamberlin, 1941:15, pl. 3, fig. 28. Alnplocheir sequoia Chamberlin, 194937, figs. 12-13. any form of M. sequoia. Finally, the solenomere Chamberlin andHoffman, 1958: 17.Buckett, 1964:7. arises subbasally from the acropodite as in M. Motyxia sequoia Causey and Tiemann, 1969:29. tularea, instead of on process "B"as in M. sequoia. Motyxia sequoia sequoia Causey andTiemann, 1969:330- 31, figs. 31-32. Motyxia sequoia (Chamberlin) Diagnosis. Prefemoral process divided basally Type specimens. Male holotype and one male into relatively narrow lateral and variably broad and one female paratypes (NMNH) collected by S. medial subbranches, former bowed strongly or gen- and D. Mulaik, 20 March 1941, 10 mi (16 km) E tly anteriad, apically relatively narrow, subequal Hammond, Tulare County. in length to, or moderately shorter than, medial Diagnosis. Medial branch of prefemoral pro- subbranch, latter often closely appressed to cess narrow and sublinear, apically acuminate; acropodite, with variable basal spur, breadth vari- solenomere relatively long, subequal in length to able, leaning or curving apically, tip broad to acumi- distal part of process "B"; latter narrow, sublinear

Figs. 21-24 (facing page). Motyxia tulal,ea ollae. 21, acropodite ofleft gonopodof holotype, medialview. 22, the same, lateralview. 23, the same, subcaudalview. 24, acropodite of lekgonopodofa male from Deer Creek Road atPothole Creek, medialview. Abbreviations as in figs. 3-8, 9-12. Scale line = 0.50 min for all figs. Figs. 25-31 (facing page). Motyxia sequoia. 25-27, M. s. sequoia. 25, left gonopod of a inale from 8 mi (12.8 km) "above" Hammond, medial view. 26, acropodite of the same, lateral view. 27, left cyphopod of female from the saine locality, caudal view. 28-29, M. s. alia. 28, acropodite of left gonopod of holotype, inedial view. 29, the same, lateral view. 30-31, M. sequoia intergrades. 30, acropodite of left gonopod ofa male from 4.1 mi (6.6 km) N Clough Cave, medial view. 31, the same, lateral view. Abbreviations as in figs. 3-8,9-12.Scale line = 1.00 mm for all figs. INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 343 344 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI for most of length, curved gently mediad or reflexed Yakohl Valley, and absent from males near Balch caudad distally, apically subacuminate (Figs. 25- Mountain Park. The chief diagnostic feature, how- 27). ever, the length of the solenomere, less than half Variation. On the prefemoral process, the the length of the distal part of process "B", does not relative lengths of the subbranches vary, although vary, nor does the position of its detachment from the lateral is generally shorter than the medial, the latter. which tends to be slightly wider and curve broadly Measurements of pinned adults are summa- mediad in southern males. The most dramatic rized in table 7; no generalizations are possible changes, however, involve the basal spur, which from these data. becomes proportionally longer 8-12 mi (12.8-19.2 Distribution. From the vicinity of Three Riv- km) south and east of Hammond such that the ers, on highway 198, to Clough Cave, in the south- process appears three-branched. On the acropodite, western corner of Sequoia National Park, to Balch the length of the solenomere relative to the distal Mountain Park. Dimensions are 16 mi (25.6 km), part of process "B" varies, although the former is northlsouth, and 11.1 mi (17.8 km), eastlwest. Spec- always at least 213 to 314 as long. Process "B" may imens were examined from the following new local- also be reflexed caudad distad or curved gently ities: Tulare Co., 10 mi (16 km) SE Three Rivers, mediad. along S Fork Kaweah R., 5M, 3 May 1979, J. T. Measurements of pinned adults are summa- Doyen (UCB); and 13mi (20.8 km) SE Three Rivers, rized in table 6; no generalizations are possible South Fork Camp, 3M, 3 May 1979, M. F. Burgler from these data. (UCB). Distribution. The vicinity of Hammond on the East Fork of the Kaweah River along California Motyxia sequoia intergrades highway 198, extending into the adjacent fringe of Figs. 30-31 Sequoia National Park. Specimens were examined from the following new locality, buried in debris in Geographically intermediate specimens west a concrete sluice high on the mountainside: Tulare and northwest of Clough Cave are also anatomical- Co., 2.8 mi (4.5 km) E Hammond, Kaweah Power- ly intermediate in the conditions of the medial house #3, M, 4F, juv., 1 April 1989, N. J. Smith, R. branch of the prefemoral process, the solenomere, D. Haines (NCSM) and MM, FF, 22 April 1991, N. and process "B". The first and last are broader than J. Smith, R. D. Haines, R. M. Shelley (NCSM). in the nominate subspecies but narrower than in M. s. alia,and the solenomere is more than half as Motyxia sequoia alia Causey and Tiemann long as the distal part of process '73" but never equal Figs. 28-29 to the latter and hence shorter than in the nominate race. Measurements of pinned adults are sum- Motyxia seqzsoia alia Causey and Tiemann, 1969:29, marized in table 8; on the average, females are figs. 5, 29-30. larger than males in both dimensions. Distribution. The area west and northwest of Type specimens. Male holotype, female allo- Clough Cave, in the southwestern corner of Se- type, and 5 male and 6 female paratypes (FSCA) quoia National Park and the adjoining part of the collected by D. L. Tiemann and D. D. Linsdale, 6 Sequoia National Forest. As Causey and Tiemann March 1964, along California highway 198at Horse- (1969) confusingly record intergrades in the sub- shoe Creek, SE Lake Kaweah, Tulare County. species accounts, I record below all samples that I Diagnosis. Medial branch of prefemoral pro- examined: Tulare Co., Sequoia Nat. Pk., 1 mi (1.6 cess variably broad to subspatulate, apically round- km) W Clough Cv. Ranger Sta., 3M, 14F, 25 March ed; solenomere short, less than half as long as distal 1967, DLT (FSCA); 4.1 mi (6.6 km) N Clough Cv., part of process "B; latter broad to subspatulate, 2M, 25 March 1967, DLT (FSCA); 4.1 mi (6.6 km) curvilinear (Figs. 28-29). NW Clough Cv., 2820f,4M, 2F, 25 March 1967, DLT Variation. The breadths of the medial prefem- (LACMNH); and 8.7 mi (13.9 km) NW Clough Cv., oral subbranch and process "Bvary, and the latter 9M, 12F, 25 March 1967, DLT (FSCA, LACMNH). may be suberect or curve slightly caudad. The most Remarks. A male from the last site has a variable structure is the prefemoral spur; it is long gonopod on the right, at the anterior position on in the types, short, broad, and trifurcate in males segment 7, and a normal walking leg on the left, a near Terminal Reservoir, very short in ones from classical example of hysterotely (Shelley 1977). INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 345

Figs. 32-35. Motyxiapior. 32, left gonopod of a male from 6.8 mi (10.9 km) S Badger, medial view. 33, acropodite of the same, lateral view. 34, thesame, caudalview. 35, leftcyphopodofafemale from thesalne locality, caudalview. Abbreviationsas infigs. 3-8,9-12. Scale line = 1.00 mm for all figs.

Figs. 36-39. Motyxiaporrecta. 36, left gonopodoflectotype, medialview. 37, acropodite ofthe same, lateralview. 38, the same, caudal view. 39, left cyphopod of female paralectotype, caudal view. Abbreviations as in figs. 3-8. 9-12. Scale line = 1.00 mm for all figs. 346 Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Figs. 40-42. Motyxia tienzanlri. 40, acropodite ofleftgonopod ofholotype, medial view. 41, the same, lateral view. 42, leftcyphopod of female allotype, caudal view. Abbreviations as in figs. 3-8. 9-12. Scale line = 1.00 mm for all figs.

Motyxia pior Chamberlin subacuminate; and process "B" may be broader Figs. 32-35 than shown and lean slightly anteriad or caudad. Additionally, there may be a single, moderately Motyxia pior Chamberlin, 1941:16. Chamberlin and long macroseta distad on the prefemur; it may or Hoffman, 1958:39. Buckett, 1964:9. Causey and may not arise from a minute papilla, and the latter Tiernann, 196925-27, figs. 13-16. can also exist without the seta. This seta is partic- Motyxia pior form secca Causey and Tiemann, 1969:27, figs. 15-16. (Invalidly proposed for a minor variant ularly prominent in males collected along Dry Creek, of pior and without standing in nomenclature.) south of Badger, and SEM photos make it look even larger. Causey and Tiemann (1969) therefore Type specimens. Female holotype (NMNH) thought it was a sclerotized structure representing collected by S. and D. Mulaik, 21 March 1941,12mi a second prefemoral branch, concluded that M. pior (19.2 km) NE Hammond, Tulare County. is polymorphic, and named this variant "secca" Diagnosis. Prefemoral process upright and because of its prevalence along Dry Creek. Howev- entire, a short, spiniform projection extending to er, the International Code of Zoological Nomencla- near level of distal extremity of solenomere; latter ture has no provision for naming variants, and the fused basally to process "B", detaching near authors did not officially propose a subspecies, so midlength, detached part minute, blunt, and indis- the name, "secca," has no standing in nomencla- tinct, much shorter than distal part of process "B"; ture. latter broad and irregularly semilunar, suberect, Measurements of pinned adults are summa- medial margin curvilinear, lateral margin subcon- rized in table 9; on the average females are larger tinuous, curving generally mediad, apically in both dimensions than males at the two sites with subacuminate; cyphopod with hirsute receptacle both sexes, and individuals tend to be longer and (Figs. 32-35). broader to the west in the range. Variation. The prefemoral process may or Distribution. According to a guide, biolumi- may not have a short basal tooth on either or both nescent millipeds have been seen inside Crystal sides; the detached part of the solenomere may be Caverns, Sequoia National Park. None has ever slightly longer than shown in figs. 32-34 and been collected there, but the site record is plausible INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 347

and conforms to the known distribution of M. pior. and 10.2 mi (16.3 lim), northlsouth. Specimens This is the northernmost known locality for biolumi- were examined from the following new localities: nescent millipeds, and the species' range extends, Kern Co., Greenhorn Mts., exact locality not speci- eastlwest, from Woodlake Valley north of Elder- fied, M, 24 July 1955, A. R. Hardy (LACMNH); wood to the vicinity of Hospital Rock in Sequoia along Western Divide Rd. at Rattlesnake Cr., 35M, National Park, a distance of some 16 mi (25.6 km). 2F, 21 February 1966, D. L. & D. A. Tiemann Specimens were examined from the following new (LACMNH); along Western Divide Rd. at Oak Flat, localities: Tulare Co., 6.8 mi (10.9 km) S Badger, 3M, 6F, 21 February 1966, D. L. & D. A. Tiemann along Dry Cr. Rd., 17M, 22F, 2 juvs., 1 April 1964, (LACMNH); Democrat Hot Spgs. turnoff, Kern R. DLT (FSCA, LACMNH); 9.3 mi (14.9 km) S Badger, Cyn., 23M, F, 27 January 1965, D. L. & P. M. along Dry Cr. Rd., 4M, 6F, juv., 31 March 1964, Tiemann (LACMNH); and 7 mi (11.2 km) SW Dem- DLT (FSCA); and 2 mi (3.2 km) "above" Three ocrat Hot Spgs., Upper Richbar Cpgd., 34M, 29F, 22 Rivers, 3M, 3F, DLT, D. D. Linsdale (FSCA). December 1967,J. S. Buckett, M. R. Gardner (UCD). Motyxia porrecta Causey and Tiemann Motyxia tiemanni Causey Figs. 36-39 Figs. 40-42

Motyzia porrecta Causey and Tiemann, 1969:27, 29, Motyxia tiel~zalrlriCausey, 1960:132-135, figs. 1-2. Cau- figs. 7, 21-22. sey and Tiemann, 1969:31-32, figs. 9, 23-24.

Type specimens. Male holotype, female allo- Type specimens. Male holotype and female type, and 6 male and one female paratypes (FSCA) allotype (AMNH), two male and two female collected by D. L. Tiemann, 27 January 1965, at paratypes (NMNH), and one male and one female Stork Creek county picnic site, Kern River Canyon, paratypes (FSCA) collected by D. L. Tiemann, 11 Kern County. Twenty-six male paratypes (FSCA) May 1959, at Shirley Meadows, ca. 6 mi (9.6 km) W and 5 male paratypes (LACMNH) talien by same north arm of Lake Isabella, Sequoia National For- collector on same date, 1.3 mi (2.1 km) "below" est, Kern County. Three male paratypes (FSCA) Miracle Hot Springs, Kern River Canyon, Kern talien by same collector in May 1959. County. Diagnosis. Prefemoral process upright and entire, very broad basally, narrowing slightly at Diagnosis. Prefemoral process entire, moder- midlength then expanding again to broad tip, slight- ately long and gently sinuate, apically rounded, ly shorter than, or subequal in length to, process extending to near level of distal extremity of sole- "B"; solenomere arising basally from acropodite, nomere; latter fused basally to process "B", detach- upright for most of length, curving slightly cau- ing proximal to midlength, detached part short but dolaterad apically and overlapping process "B"; distinct, clearly extending from process "B,apical- latter long and upright, narrowly segregated basally blunt; process "B" long and subspatulate, distal ly from solenomere, expanded basally, narrowing part curving in gently sinuate configuration but gradually distad to subacuminate tip; cyphopod extending nearly directly ventrad, sides tapering with glabrous receptacle (Figs. 40-42). gradually distad to rounded tip; cyphopod with Variation. The general configurations of the glabrous receptacle (Figs. 36-39). branches are constant, but their relative lengths Variation. The only gonopodal variation in- vary. The prefemoral process may be subequal in volves slight differences in the length and curva- length to process "B", and the solenomere varies ture of the telopodal branches. Measurements of from 112 to 213 as long as the latter. Measurements pinned adults are summarized in table 10; females of pinned adults are summarized in table 11; fe- are slightly larger in both dimensions than males at males at Shirley Meadows are larger in both dimen- two of the three sites with both sexes, but no sions than males, and individuals from east of geographic trends are evident. Glenville tend to be longer than those to the north- Distribution. The Kern River Valley from east from Shirley Meadows. below the entrance to the Sequoia National Forest Distribution. The Greenhorn Mountains of to near Miracle Hot Springs, and north to Oak Flat northern Kern County, from just below the Tulare on the Western Divide (Rancheria) Road. Approxi- County Line to Evans Flat Campground, some 9.6 mate dimensions are 20.8 mi (33.3 km), eastlwest, mi (15.4 km) to the south. Specimens were exam- 348 Volume 11, Nos. 3-4, September-December. 1997, INSECTA MUNDI ined from the following new 1ocalities:Kern Co., inoperative when an individual is placed in dark- E of Glenville, M, 18 March 1941, S. and D. Mulaik ness during normal daylight hours. (NMNH); and Alder Cr. Cpgd., ca. 2.5 mi (6.3 km) No author has suggested a function for the NW Shirley Mdws., 2M, 24 March 1970, H. B. luminescence. Davenport et al. (1952) discount Leech (CAS). recognition, because polydesmoids are blind, and correctly state that the "warning luminescence" Remarks and observations on the hypothesis can only be proved by controlled field bioluminescence experiments. However, this is still a valid hypothesis, and few others seem plausible. Luminescence A quarter-century has elapsed since the last could conceivably function as a nighttime equiva- publication on Motyxia, and a summary of current lent of aposematic coloration, as exhibited by east- knowledge of the bioluminescence is in order be- Nearctic xystodesmids and by Sierran representa- cause of the phenomenon's significance to the tives of Sigi~~ocheir(Whitehead and Shelley 1992, Diplopoda as a whole; I supplement this with per- Shelley 1995a), because the glowing millipeds are sonal observations on live M. s. sequoia and M. pior, quite conspicuous. Causey and Tiemann (1969, which I have collected and received from California 1970) report that they sometimes "resemble the entomologists. Loomis and Davenport (1950), Dav- starry sky on a dark night," and while I have not enport et al. (1952), and Causey (1960) describe the viewed this situation, I have seen several glowing luminescence as a "greenish-white" glow, but the Motyxias walking in a dish. The luminescence is a color that I observed was neon white without a spectacular phenomenon and without question greenish tint, resembling the glow from a "light would "warn" predators of the millipeds' presence. stick." The luminescence is continuous and not under voluntary control; it seems inexhaustible, Acknowledgements and captured individuals of M, s. sequoia glowed brightly for many hours on 4-5 consecutive eve- Field work associated with this study was sup- nings, until they either died or were preserved. The ported by grants 3871-88 and 4495-91 from the entire dorsum, legs, and antennae glow evenly; I National Geographic Society; laboratory examina- have not checked the ventral surface, but according tions of the specimens occurred in 1992, when I was to Causey and Tiemann (1970), the sterna and working at the NMNH on a National Science Foun- gonopods do not emit light. The reaction requires dation Mid-Career Fellowship. I thank the follow- oxygen and takes place in "deeper layers of the ing curators for loaning the indicated type and non- integument"; Davenport et al. (1952) did not ob- typical specimens under their care: J. A. Codding- serve light from egg masses of M. sequoiae or first ton (NMNH), paratypes of M. tieinal~i~i,holotypes instars individually, but masses of the latter glowed and paratypes of M. inol~ica,expansa, exilis, kerr~a, faintly. Older instars apparently did illuminate, and pior, Luinii~odesinussequoiae, and Xystocheir because the authors noted a decrease in intensity at tularea and sequoia; G. B. Edwards (FSCA), syn- the time of molting, which corresponds with a types of M. porrecta, holotype, allotype, and reduced metabolic rate. paratypes of M. sequoia ollae, holotype and Causey and Tiemann (1969, 1970) repeatedly paratypes of M. s. alia, and paratypes of M. tieinan- stated that individuals within populations differ as i~i;N. I. Platnick (AMNH), holotype and allotype of to the intensity of the light and that some are dark M. tieinan~~i;and the late C. L. Hogue (LACMNH), and do not luminesce. Similarly, I have had individ- syntypes of M. porrecta. Other non-typical speci- uals that glowed brightly immediately upon expo- mens were provided by the following curators and sure to darkness and others that required handling collection managers: W. J. Pulawski (CAS), J. A. for 15-20 minutes before they began emitting a Chemsak (UCB), the late R. 0. Schuster (UCD), faint light. Causey and Tiemann do not indicate the and S. I. Frommer (UCR). N. J. Smith and R. D. times of their observations, but mine came at differ- Haines kindly sent specimens to the NCSM and ent hours of the day, the former at night and the provided valuable field assistance in California. R. latter during daylight, and suggest a circadian L. Hoffman provided constructive commentary on rhythm. Controlled experimentation on photoperi- a preliminary draft of the manuscript; R. G. Ku- od may therefore be profitable because the reaction hler, NCSM scientific illustrator, assisted with the may observe a periodicity, taking place immediate- figures; and J. C. Beane measured the pinned ly in normal hours of darkness, but being slow or LACMNH specimens. Finally, I would be remiss INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 349

not to acknowledge the outstanding collecting ef- Chamberlin, R. V. 1947. Some records and descrip- forts of Darwin L. Tiemann, who secured more tions of diplopods chiefly in the collection of the bioluminescent xystodesmids than all other collec- Academy. Proceedings of the Academy of Natural tors combined. Sciences, Philadelphia, 99:21-58. Chamberlin, R. V. 1949. Some millipeds of the families References Polydesmidae and Xystodesmidae. Journal of the Washington Academy of Science, 39:94-102. Attems, C. G. 1938. Polydesmoidea 11. Fam. Leptodesmi- Chamberlin, R. V., and R. L. Hoffman. 1958. Check- dae, Platyrhacidae, Oxydesmidae, Gomphodesmi- list of the millipeds of North America. United States dae. Das Tierreich, Lief 70:l-576. National Museum Bulletin No. 212: 1-236. Bollman, C. H. 1893. The Myriapoda of North America. Cook, 0.F. 1904. Myriapoda of northwestern North United States National Museum Bulletin No. 46:l- America. Iiz: Harriman Alaska Expedition, 8(In- 210. sects, pt. 1):47-82. Buckett, J. S. 1964. Annotated list of the Diplopoda of Davenport, D., D. M. Wootton, and J. E. Cushing. California. Simmons Publ. Co., Davis, California, 1952. The biology of the Sierra luminous millipede, Lulninodesinus sequoiae, Loomis and Davenport. 34 PP. Buckett, J. S., and M. R. Gardner. 1968a. Revision of Biological Bulletin, 102:100-110. the milliped genus Harpaphe Cook fi-om western Hoffman, R. L. 1964. A new subfamily of xystodesmid North America (Polydesmida: Xystodesrnidae). Cal- millipedes from North America and China (Poly- ifornia Department of Agriculture Bureau of Ento- desmida). Transactions of the American Entomo- mology, Occasional Paper No. 11: 1-51. logical Society, 90:301-311. Buckett, J. S., and M. R. Gardner. 1968b. Revision of Hoffman, R. L. 1980 ("1979). Classificatioiz of the the milliped genus Wainolzia Chamberlin from the Diplopoda. Museum &Histoire Naturelle, Geneva, Sierra Nevada of central California (Diplopoda: Switzerland, 237 pp. Polydesmida: Xystodesmidae). Proceedings of the Jeekel, C. A. W. 1971. Nomenclator generum et famil- Biological Society of Washington, 81:511-538. iarum Diplopodorum: A list of the genus and family- Buckett, J. S., and M. R. Gardner. 1969a. A new group names in the class Diplopoda from the 10th genus of xystodesmid milliped from northern Cali- edition of Linnaeus, 1758, to the end of 1957. fornia. Entomological News, 80:67-73. Monografieen van de Nederlandse Entomologische Buckett, J. S., and M. R. Gardner. 1969b. A new Vereniging. No. 4:l-412. species of Aizolnbrocheir Buckett and Gardner from Loomis, H. F. 1953. New millipeds of the western states the inner Coast Ranges of northern California. and lower California. Journal of the Washington Entomological News, 80:293-299. Academy of Science, 43:417-422. Buckett, J. S., and M. R. Gardner. 1969c. A new Loomis, H. F., and D. Davenport. 1951. A lumines- genus of xystodesmid milliped with the description cent new xystodesmid milliped from California. of a new species from Idaho (Diplopoda: Xystodes- Journal of the Washington Academy of Science, midae). Proceedings of the Entomological Society of 41:270-272. Washington, 71:65-70. Shelley, R. M. 1977. Appendicular abnormalities in the Causey, N. B. 1955. New records and descriptions of milliped family Xystodesmidae (Polydesmida). Ca- Californian Diplopoda. Proceedings of the Biologi- nadian Journal of Zoology, 55: 1014-1018. cal Society of Washington, 68:87-94. Shelley, R. M. 1987. The milliped Steizodeslnus tuobi- Causey, N. B. 1960. A third luminous milliped, Motyxia tus (Chamberlin) (Polydesmida: Xystodesmidae) in tiemalzlzi, n. sp. (Xystodesrnidae: Polydesmida). Texas and New Mexico. National Geographic Re- Wasmann Journal of Biology, 18:131-135. search, 3:336-342. Causey, N. B., and D. L. Tiemann. 1969. A revision of Shelley, R. M. 1989. Rhysodesinus chisosi, n. sp., a the bioluminescent millipedes of the genus Motyxia biogeographically significant milliped from the Chi- (Xystodesmidae: Polydesmida). Proceedings of the sos Mountains, Texas (Polydesmida: Xystodesmi- American Philosophical Society, 113: 14-33. dae). Southwestern Naturalist, 34:219-224. Causey, N. B., and D. L. Tiemann. 1970 ("1969"). The Shelley, R. M. 1992. Occurrence of the milliped, Sten- bioluminescent millipedes of the genus Motyxia. odesinus tuobitz~s(Chamberlin), west of the Rio Bulletin du Museum National &Histoire Naturelle, Grande (Polydesmida: Xystodesrnidae). Insecta 2e Serie, 41(2):35-39. Mundi, 6:19-21. Chamberlin, R. V. 1941. New western millipeds. Bul- Shelley, R. M. 1993a. The milliped genus Orophe Cham- letin of the University of Utah 31(12)[Biological berlin (Polydesmida: Xystodesrnidae). Insecta Mun- Series 6(5)]:1-23. di, 7:175-182. Chamberlin, R. V. 1944. Two millipeds fi-om southern Shelley, R. M. 199310. A new xystodesmid milliped California. Proceedings of the Biological Society of genus and species from Oregon and Washington Washington, 57: 113-116. (Polydesmida). Myriapodologica, 2:91-100. Volume 11, Nos. 3-4, September-December, 1997, INSECTA MUNDI

Shelley, R. M. 1993c. The milliped genus Isaphe Cook Tiernann, D. L. 1964. Grant No. 3424 - Penrose Fund (Polydesmida: Xystodesmidae). Canadian Journal (1963), $880. The distribution of luminous species of Zoology, 71: 1161-1168. of millipeds in California. Year Book of The Amer- Shelley, R. M. 1994. The Chonaphini, a biogeographi- ican Philosophical Society, 1964:298-299. cally significant milliped tribe in eastern and west- Verhoeff, K. W. 1941. Uber Gruppen der Leptodesmiden ern North America. Brimleyana, 20: 111-200. und neues System der Ordo Polydesmoidea. Archiv Shelley, R. M. 1995a. The Sigmocheirini, a xystodesmid fur Naturgeschichte, 10:399-415. milliped tribe in the Sierra Nevada Mountains, Whitehead, D. R., and R. M. Shelley. 1992. Mimicry California, U. S. A. (Polydesmida: Xystodesmidae). among aposematic Appalachian xystodesmid milli- Entomologica Scandinavica, 26:339-360. peds (Polydesmida: Chelodesmidea). Proceedings of Shelley, R. M. 199513. Parcipro~nr~s,n. gen., a xys- the Entomological Society of Washington, 92:177- todesmid milliped genus in the Sierra Nevada Moun- 188. tains, California (Polydesmida). Myriapodologica, Wood, H. C. 1867. Notes on a collection of California 3:53-70. MYRIAPODA, with the descriptions of new eastern Shelley, R. M. 1996. Revision of the milliped genus species. Proceedings of the Academy of Natural Xystocheir Cook (Polydesmida: Xystodesmidae). Sciences, Philadelphia, Canadian Journal of Zoology, 74:1336-1363. 19:127- 130. Tiemann, D. L. 1963. Grant No. 3224 - Penrose Fund (1962), $500. Investigations of the distribution of luminous species of millipeds in California. Year Book of The American Philosophical Society, 1963:353-354.

Tables 1-11.

Table 1. hltsrurrmldunl m

rn muwrmml wit.mun mnrwimm) \~l,r.rillia rn mnr w tmml wit.muo marwrmm) wil.rsllo Krm Ca.. Kcrnr Tulsre Co., Coy Flnl N.Camp N~lran

Kern Co., Fort Telon Tulwe Co., Camp Bcmnp

Lo. ~ngalcrco.. ~opmp.Canyon Rd. st Crccnlcar cpnyon Tulsra Co.. along CA 11~.190 slt ~oorrhousecr.

Table 2. hlcasur

Mabs n - 10 Ftmnles 28.26 5.59 19.78 31.45 6.50 20.67 mnge 26.60-30.14 5.03.6.20 18.13-21.23

Tulnrc Co.. Dwr Creek Rd.. Tulsrt Ca.. CmeRhorn MI. .. 0.9 ml (1.4 km) N Kern Co. Une

Tul-e CO.. 2 ml (32 km) S Plna Flut. ceptnrro Cr.

Tulnrt Co., 3.3 ml (5.3 kml SE Whitr Rivrt Kern Co.. 0.2 mt (0.3 km) S Tvlnrt Co. llns

i~lnlesn = 28 Fcmacs n = 4 24.22 4.72 19.47 26.23 5.15 19.66 nner 22.47-31.90 1.28-6.44 17.60.22.34 ran81 24.56-29.05 4.90-5.50 18.93-20.60 Tulnre Co.. 7.3 mt (11.7 km) NE WNlr mvrr INSECTA MUNDI, Vol. 11, Nos. 3-4, September-December, 1997 351

~nblr9. ~earurrmcntdstn or pinned sddcs ar M. pior, rummnriei of lndluldus~mrssurem~nt~.

merwrmml wiLlsaa m rnarlvinun) w,Lretro u!!Yl9 msr W lmnll \YILrstlo mar wrmml WILI.(lo

Tulnra Ca.. Enwanre lo Tule lndlnn Rrs. Tulnra Co., Sequal. NII. Pt.. PoMshn TmUcr Camp

Talnrr Co.. 4.l mt (6.6 km)E cnbnncr Tulr R. lndlnn Rrr Tulnrt Co.. Srguols Nnl. Pt.. tlospltd Rock Plcae Ares

Male hl~lcln = 18 Fcntsllr n = I2 29.88 5.61 18.78 30.57 5.60 18.33 33.65 6.12 19.06 NIIY;~ 26.0533.96 5.17-6.13 16.82.20.00 rnngt 30.30.36.55 6.00-6.77 17.78-20.27 Tulnrc Co.. Dcrr Cr. Rd. nt PoIholc Cr. Tulnrc Co., 0.2 mf (0.3 km) E Thrrt Rben

hlvlrr n = 2 32.00 5.20 16.26 nngt 31.24-32.75 5.08-5.31 15.51-17.00

Tulnre ca.. Dry Crrrt hlsbsn- 111 F~~PIC'n - 31 28.91 5.56 19.24 31.57 6.30 19.96 nng 23.31-34.83 4.19-6.56 16.53-22.68 rsngc 23.40-38.57 4.66-7.45 18.08-12.65

Tnblc 10. hl~srurmtmtdscr ofplmrd P~UIIS01 Of. ~01m0,1m8msui~~ 01 lndlvldusl mr%xuremmls m nnnr W rmm) \YILI.IID mar \\'(nun) mar W t-8) \YILISU. mnx wrnunl w,l.mtlo Tulue Co. 8.5m1 (13.6) E Hsnunand Ktrn Co.. Drmncrvt llot Springs Malts n = I9 Fln,dar n = 4 32.44 6.10 18.83 35.02 6.90 19.71 nnge 27.08-36.69 4.54-6.73 16.77-20.17 nngc 32.99-37.39 6.35-7.26 19.13-20.67

Tulnrr Ca.. 5.8 ml (9.3 lon) E llsmmond Kern Co.. 1.3 ml (2.1 km) W hllnrlr llal Springs

Tulnre Co.. 3.1 nd (5.0 knl) S Iinnunond Kern Co.. Wcslrrn Dlvlda Rd.. Rsr(lasn-a Cr. Mnlrx n - 32 f emelcs n = 22 32.47 6.28 19.41 32.26 6.44 20.03 nl*le$" * 34 Ftmdr, n - 3 lunge 27.26.38.41 5.40.7.19 17.37-21.38 nngc 15.68.37.54 5.15-7.40 17.88-21.93 16.03 5.24 20.14 28.87 5.63 19.51 rang< 13.18-28.52 4.33-5.99 17.62-22.14 lmngr 27.86-30.20 5.49.5.75 18.68-20.13

Kern co.. Wextrrn Dlrldc Rd.. 0- flat

hlslts 0 13 f cmslcs 0 = 6 29.99 5.80 19.37 30.40 6.08 20.06 m.rwmun1 \YILISU. msrwrmm, WiLInllo range 17.55.31.33 5.50-6.06 18.82-19.96 mnsr 28.63-31.99 5.44-6.44 17.01-21.75

Tulnrt Co., 2 ml (3.2 km) SW Thrra Rlvcn Kern Co.. urnR. Cpnyon. Slork Cr. Cnmpgrovnd

Mslrr n = 6 Femdt. n = 4 26.01 5.35 20.57 29.59 5.73 19.36 rsnea 14.00.27.21 5.11.5.84 19.64-22.00 m% 25.21-32.69 4.80.6.26 19.04-19.83

Tulnre Ca., hfllo

Tulnrt Co.. apporltrTrmind Relrrvot mnr Wlmml WiLIpUo m n>er\v rmm, w,Lretro

tiem Co., Grrrnhorn hU%.,Shtrlry Me.dorr

Tulnrt Co.. Iml(1.6 km) NW Clough Cavt. Sequals N.1. PL

Krrn Co., 8.3 ml (13.3 lon) E ClmvUb

M~hln = I8 30.81 5.36 17.39 nng 23.83-39.87 4.86-5.78 13.91-21.63

T~bls8. hlsslur~mcntdate or pinntd ldullr of .*I. rq#qrofn Inlergnder. summnrirs ol indlrldud mcslurrm~nll.

msx Wrmml m mnrWlmml wl.rstlo

Tulnrt Co., 4.1 nd (6.6 Um) NW Clou&h Cnvr

hlslr. n = 4 F

Tulnra co.. 8.7 ml (13.9 km) NWclovgh care

h1.h~ n = 10 Fcm~Iesn - 10 29.58 5.55 18.78 34.99 6.54 18.74 rwrgt 27.35.32.22 4.84-6.11 17.38-20.19 mga 31.69-39.02 5.83-6.86 17.40.20.74