The Milliped Genus Bollmanella (Diplopoda, Choreumida, Conotylidae)

Even while my recent (97o) revision of the milliped Family Conotylidae was in press, new data on the family had begun to accumulate. This paper is the first of a series of supplemental reports designed to update our knowledge of the conotylids, a group of millipeds of considerable importance in biogeography. Like many other milliped genera and species, the genus Bollmanella, and its single species B. oregona, have remained enigmatic since they were described from a single male specimen by R. V. Chamberlin in I941. The description of the genus alluded mostly to body form and color, and even contradicted the specific diagnosis of the only included species. The description of B. oregona also contained errors, and no illustrations were provided. As if this were not enough, the type locality suggested an error of several hundred miles. The type of B. oregona was in the Chamberlin collection in Salt Lake City. The diplopod portion of the collection has remained uncurated since Chamberlin’s death and is in a confused state, but while my conotylid revision was in press, Mr. Thomas Lorenz, then in charge of the collection, found the holotype of B. oregona and loaned it to me. In I973, I was fortunate in receiving a large number of Berlese extraction samples from Mrs. Ellen Benedict, Portland State University, Portland, Oregon, and from Dr. David Malcolm, Pacific University, Forest Grove, Oregon. These samples were rich in millipeds of many groups previously known only from a very few specimens, and included literally hundreds of individuals of the related genera Taiyutyla and Bolhnanella. Members of these genera must be among the more common humus animals in the area of coastal Oregon. As I mentioned earlier in reporting on part of these collections (Shear, I973), northern California and the state of Washington are much in need of the kind of thorough exploration by

enough, the type locality suggested an error of several hundred miles.
The type of B. oregona was in the Chamberlin collection in Salt Lake City. The diplopod portion of the collection has remained uncurated since Chamberlin's death and is in a confused state, but while my conotylid revision was in press, Mr. Thomas Lorenz, then in charge of the collection, found the holotype of B. oregona and loaned it to me. In I973, I was fortunate in receiving a large number of Berlese extraction samples from Mrs. Ellen Benedict, Portland State University, Portland, Oregon, and from Dr. David Malcolm, Pacific University, Forest Grove, Oregon. These samples were rich in millipeds of many groups previously known only from a very few specimens, and included literally hundreds of individuals of the related genera Taiyutyla and Bolhnanella. Members of these genera must be among the more common humus animals in the area of coastal Oregon. As I mentioned earlier in reporting on part of these collections (Shear,I973), northern California and the state of Washington are much in need of the kind of thorough exploration by Part of the work for this paper was done while a Richmond Fellow at Harvard University.
Manuscript received by the Editor February 6, 1974. I34

Shear Milliped Genus Bollmanella
Berlese extraction which Benedict and Malcolm have carried out in Oregon.
It is now possible to do a number of things about Bollmanella, which in 97o I had to treat as a nomen dubium. ) The genus can be characterized as fully distinct and valid, separate from, but related to, Taiyutyla and Conotyla, and more distant from dustrotyla and /lchemenides, 2) some clear ideas of the distribution of the genus can be gained, and 3) seven new species can be described.
The distribution of the eight known species of Bollmanella ranges from southern coastal Oregon to just above the Columbia River in Mason Co., Washington, with one species reported here from the Vgallowa Mountains in extreme northeastern Oregon. The scattered nature of the localities suggests that species of Bollmanella, except for B. oregona, have small ranges, and we might expect many more to be discovered. In particular, the isolated mountain ranges of eastern Oregon need exploration.
Ecologically, Bollmanella species have been collected exclusively from litter and duff derived from deciduous trees. Conifer duff and litter support species of Taiyutyla, though members of this latter genus are also found in deciduous duff, often syntopically with Bollmanella species. Although B. oregona ranges from sea level to. around 2ooo' elevation, most of the other species seem to have been collected from forests or parkland between 900' and I5OO' elevation. This contrasts again with some species of Taiyutyla, which (unpublished data) are known from quite high elevations. Species of an unrelated chordeumid genus, Rhiscosomides (Shear,I973), are found at low elevations and seem to favor coniferous forests. This ecological data also represents information from many collections which did not contain specimens of Bollmanella, thus clearly suggesting which habitats are most favorable.
All shaped process between the coxae in most species. Third and fourth legpairs of males enlarged, strongly bowed, but with, at most, small mesal femoral knobs. Legpairs 5-7 either normal in size and form, or decreasing from slightly enlarged to normal in size. Prefemora of male legpair with mesobasal processes. Female genitalia without any useful taxonomic characters.
Remarks: Bollmanella is clearly related to Taiyutyla. with which it is entirely sympatric, and often syntopic. However, the uniformly smaller size, the flagellar branch of the posterior gonopods, and the T-shaped process of the posterior gonopod sternum clearly set Bollmanella species apart as a distinct phyletic line.
I would like to comment further on the Family Conotylidae as a whole, but several factors preclude it at this time. The genus Taiyutyla remains to be revised, and is affected by, as are any conclusions about Conotylidae at this time, the description by Loomis and Schmitt (I97I) of several extremely interesting new higher taxa related to conotylids, including new subfamilies and a new family.
The Taiyutyla material I have examined so far will also enable me, in this projected future paper, to comment in detail on the functional aspects of the gonopod complex. I am still unable to find any taxonomically useful characters that will allow reliable separation of females of the various species of conotylids, and members of Bollmanella are no exception. However, in the hope that some usable characters might turn up in the future, I have designated bona fide female paratypes where possible. I do not present a key to the eight known species, as they can readily be separated by reference to the illustrations of the gonopods. The gonopods of these species are so small they must be mounted on slides temporarily (in glycerine) to see detail.  There is no John Day Creek in Douglas Co., but there is a town named Days Creek, and a stream ot that name, a tributary ot the South Umpqua River. Probably this is the type locality, and not the John Day River ot Oregon's semiarid northeast.
Females similar to males, slightly larger and more robust. Notes: Chamberlin's brief discussion of the leg modifications is confusing. In the generic diagnosis leg 5 is described as being "normal" while in the specific diagnosis, leg 5 is described as "less thickened, (the) fourth joint with a subconical process on mesal side toward proximal end ." Thus B. oregona is immediately excluded from the very genus it is being described under! Chamberlin saw 14 ocelli on each side, and gave the size as 7-7.5 mm. The male holotype is in poor condition.
Where ecological data is available, the species appears to occur in duff rom deciduous trees. Many samples of coniferous duff did not yield Bollmanella oregona. The species has been collected at elevations trom sea level to I9OO t, and thus appears to be primarily a coastal or river valley species, since many samples rom higher elevations did not contain B. oregona. This is the most widespread and common species of Bollmanella. Bollmanella reducta n. sp.
Description" Male holotype. Length, 5.o ram. Ocelli 8 on each side of head. Legpair 3 much as described for B. oregona. Legpair 4 less enlarged, lacking basal femoral knobs present in B. oregona. Legpairs 5-7 of normal size. Anterior gonopods (Fig. 3) much as in oregona, but gonopod tip blunter. Posterior gonopod colpocoxites ( Fig. 4) of reduced complexity when compared to following species; flagellum () short; sheathing structure (s) fused anteriorly to body of colopocoxite, which is broad and laminate, with minute teeth distally. T-shaped process of sternum present, but broken in making temporary slide of gonopods. Pigmentation light.
Distribution: Known only from the type locality. The holotype came from a Berlese sample of Oregon oak and buckbrush litter on a steep slope.
Female similar to male in nonsexual characters, somewhat more robust and larger.
Distribution: Knovn only 1:rom the type locality. The types came 1:tom a sample o1: vine maple and Douglas fir duff. Bollmanella unca n. sp.
T-shaped process rom posterior gonopod sternum much the largest in genus, arms extending laterad (Fig. o). Pigmentation typical, but somewhat darker than in B. oregona.
Mature females not collected. Description: Holotype male. Length,9.2 mm. 2o ocelli on left side of head, 21 on right side. Legpairs 3 and 4 enlarged, approximately same size, both with small mesal femoral knobs. Legpairs 5-7 decreasing in size, legpair 7 normal in size and form. Anterior gonopods (Fig. II) upright, curved, with small lateral spine and large blunt mesal branch, presenting bifurcate appearance with telopodite. Posterior gonopod colpocoxites (Fig. I2) rather smaller than usual, flagellum not observed, probably concealed by large sheath (s) which is free from coxite for most of its length. Body of coxite with large, spirally curved basal branch (b) bearing near its base a curved lamina. Sternal process (T) much less prominent than in other species. Pigmentation very light.
Females as usual, similar in general appearance to males, but slightly larger and more robust.
Distribution: Known only from the type locality.
Note: This species is far removed from the other members of the genus, being found in the foothills of the Wallowa Mountains in extreme northeastern Oregon. This part of the state has not been much explored by biologists, and other species of Bolhnanella may well ocur in other isolated mountain ranges. The region is also close to that studied by Loomis and Schmitt (1971), and in which they found several highly unusual chordeumid species. Bollmanella bifurcata shows some intermediacy in size, pregonopodal leg modifications, and gonopod form between species of Bollmanella and Taiyutyla, and this may suggest that the two genera could be difficult to keep separate in eastern Oregon. Loomis and Schmitt (97)  Description: Male holotype. Length, 6.o mm. Ocelli 7 on both sides of head. Femora o1: legpair three very much enlarged and bearing small knobs mesally just above inflection point o1: curve. Legpair 4 somewhat smaller, 1:emora with large capitate knobs just distad of midpoint on mesal side. Legpairs 5-7 only slightly larger than norreal. Anterior gonopods (Fig. I3) much as usual, curved laterad, acuminate. Posterior gonopod colpocoxites (Fig. 4) somewhat larger than in other species, flagellum ([), long, curved into sigmoid sheath (s) body of coxite with mesal fimbriate ridge, apically laciniate. Sternal process (T) small. Pigmentation dark.
Mature females definitely belonging to this species were not collected.
Distribution" In addition to the type locality, a single 1:emale from 6 mi east ot: Allegany on the Millicoma Tree Farm may also be this species.
Notes: The type was taken in a Berlese sample o1: myrtle and Rhododendron litter and duff; the second 1:emale re1:erred to above also came from myrtle litter. Bollmanella camassia n. sp.
Description" Male holotype. Length, 7.0 ram. Ocelli I2 on both sides of head. Legpair 3 much less enlarged than in other species, with a small knob near distal ends of femora, which are evenly curved and swollen. Legpair 4 only slightly larger than normal, femoral knobs basal in position. Legpairs 5-7 normal in size and form. Anterior gonopods (Figs. 15 I6) simple, acuminate, curved in lateral view, gland channel (?) originates in deep anteriolateral depression. Posterior gonopod colpocoxites (Fig. 7) with long flagellum ([) bearing irregular membranous process near base; sheath (s) arising from midpoint of coxite; coxite bears a strong decurved process apically and a small basal branch (b). Sternal process not observed, perhaps absent. Pigmentation as in B. oregona.