taxonID	type	description	language	source
C2099450FFEC8B5337C2FBA9FB1CF93E.taxon	description	Subfamily Ochrogrammatinae Shear and Marek, new subfamily	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFEC8B5337C2FBA9FB1CF93E.taxon	type_taxon	Type genus. Ochrogramma Gardner and Shelley, 1989	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFEC8B5337C2FBA9FB1CF93E.taxon	diagnosis	Diagnosis. Distinctive in the modification of the mandibular stipes of males, which is extended ventrally into a rounded, bordered flange (mf, Fig. 13, 14, 25), and in the extraordinary length of the male gonapophyses (g, Fig. 2, 15), which are as long as or longer than the second leg telopodites. The first legpair (Fig. 1) is short, and the postfemora and tibia mesally carry long, slightly twisted setae. The second legpair coxae are extended into long gonapophyses that are longer than the reduced telopodites of those legs; the openings of the vas deferens are at the bases of the gonapophyses (vd, Fig. 15). The coxae of the third legpair (cx 3, Fig. 2, 16) have large, flattened lobes; the reduced telopodites (t 3, Fig. 2) are articulated laterally and distally on these lobes. The gonopods of the males bear on various surfaces complicated, small, much-branched tree-like (arborescent) processes (i. e., Fig. 4, 7, 8, 21), and some have a reduced, single-articled telopodite (i. e., t, Fig. 5). The color of freshly collected material is dark brownish or purplish gray, with a distinct yellow median dorsal stripe that may be less prominent anteriorly and posteriorly. This combination of characters is not found in caseyid genera other than Ochrogramma and Vasingtona. Included genera. Ochrogramma Gardner and Shelley, 1989 and Vasingtona Chamberlin, 1941.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFEC8B523580F94FFDABFEBB.taxon	type_taxon	Type species. Caseya fasciata Chamberlin, 1941, by original designation. Notes. Vasingtona consists of a single species; V. fasciata being a junior synonym of V. irritans Chamberlin, 1910, so our comments below apply to the genus. Additional information, including a diagnosis and detailed description, is in Gardner and Shelley (1989).	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFED8B5F3580FECCFC2BFAC3.taxon	description	(Fig. 1 – 12)	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFED8B5F3580FECCFC2BFAC3.taxon	description	In addition to the mandibular flanges, the males of V. irritans exhibit a typical ochrogrammatine suite of secondary sexual modifications of the anterior legs, as described in the subfamily diagnosis, in addition to which the seventh legpair coxae have posteriorly projecting lobes (cx 7, Fig. 3). The gonopods of V. irritans (Fig. 4 – 8) superficially appear very complex, due in large part to the great elaboration of the highly branched arborescent processes characteristic of the subfamily. Gardner and Shelley (1989) applied a then-current terminology to the various parts of the gonopod that in our opinion conceals homologies with Ochrogramma gonopods and those of other caseyids and striarioids. We propose a more general terminology based on studies of the gonopods of many families and genera of chordeumatids (Shear and Leonard 2007). In anterior view (Fig. 4) the gonopod sternum (s, Fig. 4) can be seen to be divided in the midline by a suture spanned with a poorly sclerotized membrane. The bodies of the coxae (c, Fig. 4) are reduced to curved, flattened plates from which arise the massive anterior angiocoxites (aac, Fig. 4), their anterior surfaces completely covered with arborescent processes (Fig. 7); mesally the anterior angiocoxites are attenuated into long processes sharply curved at their tips. In lateral view (Fig. 6), the posterior angiocoxites (pac, Fig. 6), likewise with many arborescent processes, arise basally. The poorly sclerotized colpocoxites collapse (cc, Fig. 5, 6) during SEM preparation. In posterior view (Fig. 5), the large telopodites (t, Fig. 5, 6) extend through roughly a right angle from their articulations with the coxae and there is a large, arborescent process a little below midway in their length. Because of their obvious articulation with the coxae, we consider these structures to be telopodites, but the presence of telopodites of the gonopods is otherwise rare among the Chordeumatida. A few very thin branches can be seen emerging near the curved tips of the anterior angiocoxites and these may represent flagella or flagellocoxites. The arborescent processes (Fig. 7, 8) are complex. At high magnification (Fig. 8), the terminal branches or buds flare at their tips and seem to be membranous, somewhat like tiny suction cups. What function these processes serve remains obscure. The ninth legs (Fig. 9 – 11) are typical of species of caseyids, with simple coxal processes (cp, Fig. 9, 10, 11) and single, flattened, button-like telopodite articles (t 9, Fig. 9 – 11). Near the base of each coxal process is a small pore (p, Fig. 9, 11), probably homologous to the much larger pores on the coxae of the tenth legpair. The telopodite article appears poorly sclerotized on its medial surface, where it is more densely setose (Fig. 9). One small deficiency of the otherwise excellent monograph by Gardner and Shelley (1989) is their lack of descriptions and illustrations of the tenth legpair for many of the caseyid species they describe. As seen in our illustration (Fig. 12) the coxae of these legs are enlarged, quadrate and with broad openings of the coxal glands (cxg) on their anterior surfaces. The trochanters (tr 10) appear to be at least partially fused to the coxae and bear two rounded processes mesally, a short proximal one and much longer distal one. Not surprisingly for a millipede species that ranges broadly, Gardner and Shelley (1989) noted some variation in the gonopods, as did we. However, we concur that this variation is not of a nature that would suggest splitting up the species. Since V. irritans is common and easy to collect, future work on the phylogenomics of the species should resolve questions of the significance of this variation. New records OREGON: Columbia Co.: Keystone Creek, Clatskanie Valley, N 40 ° 04.933 ′, W 123 ° 09.411 ′, 132 ′ asl, 8 February 2004, W. Leonard, C. Richart, m, f. WASHINGTON: Cowlitz Co.: Germany Creek, 3.1 mi N on SR 14, 11 November 2004, W. Leonard, ff; Germany Creek at end of county road, 5.5 mi N of Stella, N 46 ° 15.561 ′, W 123 ° 08.067 ′, 450 ′ asl, 22 November 2003, W. Leonard, C. Richart, mm; Pin Creek, Kool Road, 0.4 mi W of FishPond Road, N 47 ° 04.450 ′, W 122 ° 50.818 ′, 180 ′ asl, 25 January 2004, C. Richart, W. Leonard, ff. Grays Harbor Co.: Inner Creek at Quinalt Lake Loop Road, N 47 ° 30.15 ′, W 123 ° 46.93 ′, 13 February 2005, W. Leonard, C. Richart, f; middle fork of Satsop River, 2.9 mi N of SR 12, N 47 ° 01.98 ′, W 123 ° 31.55 ′, 31 December 2004, W. Leonard, ff; Canyon River, N 47 ° 18.210 ′, W 123 ° 30.512 ′, 11 October 2002, W. Leonard, m; N 47 ° 18.846, W 123 ° 32.474, 980 ′ asl, 13 October 2003, W. Leonard, m; 6 mi W, 1.25 mi N of Matlock, 17 January 2003, W. Leonard et al., mm, ff; Middle Fork Satsop River, 3.2 mi N of Brady, 3 January 2003, W. Leonard, C. Richart, mm; Wynochee River Road, 10 mi N of SR 12, east bank of Wynochee River, 3 January 2003, W. Leonard, m. Jefferson Co.: 12 mi E of US 101 on road to Queets Campground, N 47 ° 36.386 ′, W 124 ° 02.488 ′, 5 March 2005, W. Leonard, C. Richart, m. Lewis Co.: FR # 25 8.5 mi S of Randall, Gifford Pinchot National Forest, N 46 ° 26.456 ′, W 121 ° 59.796 ′, 1050 ′ asl, 6 December 2003, W. Leonard, C. Richart, mm, f; Stillman Basin, 1.8 mi on Weyerhauser 4000 from Dee Ell McDonald Road, N 46 ° 30.87 ′, W 123 ° 12.09 ′, 1004 ′ asl, 4 December 2004, W. Leonard, C. Richart, mm; Iron Creek Campground, 9.6 mi S of Randall on SR 25, Gifford Pinchot National Forest, 950 ′ asl, 21 December 2003, W. Leonard, f; Fisher Creek at Tilton Road, N 46 ° 35.96. W 122 ° 22.04 ′ 973 ′ asl, 21 September 2003, W. Leonard, m with spermatophore; 604 Roswell Road, Centralia, 5 January 2003, C. Richart, m; 8.5 mi S of Randall on FS 25, Gifford Pinchot National Forest, W 46 ° 26.489 ′, W 121 ° 59.890 ′, 21 December 2003, W. Leonard, m. Pacific Co.: 1.1 mi S of SR 6 on Trap Creek Road / B-line, N 46 ° 32.591 ′, W 123 ° 36.908 ′, 19 November 2005, W. Leonard, C. Richart, m, f; Ellsworth Creek Preserve, N 46 ° 23.887, N 123 ° 53.391 ′, 50 ′ asl, 23 November 2003, W. Leonard and others, mm, ff; The Nature Conservancy Preserve, 8 mi E, 5 mi N of Long Beach, 14 March 2003, W. Leonard et al, m. Thurston Co.: Ranier Road, 3.0 mi NW of Military Road, Fort Lewis Military Reservation, 29 November 2003, W. Leonard, m; Capitol State Forest, unnamed tributary to Perry Creek, N 47 ° 01.573 ′, W 123 ° 04.154 ′, 950 ′ asl, 27 October 2003, W. Leonard, f; Skookumchuck River, 5 mi S, 3 mi E of Vail, 2 October 2003, W. Leonard, mm, ff; McAllister Springs, N 47 ° 02.387, W 122 ° 43.722 ′ 60 - 123 ′ asl, 7 February 2004, W. Leonard, C. Richart, m, f; Hospital Creek, above confluence with Skoocumchuck River, N 46 ° 46.396 ′, W 122 ° 35.133 ′, 15 December 2003, W. Leonard, K. McAllister, ff; Evergreen State College, Olympia, 26 January 2003, W. Leonard, m. Wahakiakum Co.: SR 4 at milepost 23.7, 19 September 2003, W. Leonard, f; Swede Park, residence at 369 Loop Road, Grays River, N 46 ° 18.499 ′, W 123 ° 40.128 ′, 50 ′ asl, 14 January 2004, W. Leonard et al., f.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFE08B5E3580FAC3FE95FCE1.taxon	type_taxon	Type species. Ochrogramma formulosa Gardner and Shelley, 1989, by original designation. Notes. Ochrogramma is superficially similar to Vasingtona. Males of the species of Ochrogramma are usually smaller (10 – 15 mm) than those of Vasingtona irritans (16 – 22 mm). In our new interpretation of the gonopods in this subfamily, Ochrogramma is distinct from Vasingtona in having many fewer arborescent processes on the gonopod (Fig. 19, 23, 24) — two species of Ochrogramma may have none — in the reduced and perhaps sometimes absent telopodite (t, Fig. 20), having a rodlike posterior angiocoxite (pac, Fig. 20), and a prominent posterior lobe of the colpocoxite. The hypothesis that the colpocoxite originates from the coxal gland of the eighth legpair is supported by the relatively poorly sclerotized colpocoxites emerging from distinct sockets in this genus (cc, Fig. 20). Gonopod structure is similar among the species, but two of them, O. formulosa Gardner and Shelley, 1989, and O. heterogona Gardner and Shelley, 1989, seem to lack the arborescent processes (“ spicules ” of Gardner and Shelley) on the anterior surfaces of the anterior angiocoxites. We have not seen specimens of these species. Ochrogramma bentona has the most complex gonopods in the genus while those of O. haigi, O. formulosa and O. heterogona are progressively simpler. Gardner and Shelley (1989) did not give length measurements for females, but we found that females are typically longer than males (17 – 19 mm in O. bentona). The dorsal yellow stripe typical of the subfamily may be suppressed entirely in Ochrogramma species males, or present only on posterior rings. The gonopod telopodite is seen in only two of the species, O. bentona and O. formulosa. It is highly reduced and rodlike (t, Fig. 20), articulating about one-third of the way distal on the anterior angiocoxite and terminating in several filaments which are likely modified setae. The posterior angiocoxite, evidently missing in O. heterogona, is roughly similar in form to the telopodite (pac, Fig. 21, 22). Flagellocoxites are present in all species and are filamentous and branched (f, Fig. 21). Ochrogramma was considered by Gardner and Shelley (1989) to be distributed entirely to the south of the Columbia River, but this was an artifact of insufficient collecting in Washington. Ochrogramma bentona (Chamberlin, 1952), extends north of the river to the Puget Sound region and Ochrogramma skamania n. sp. is from west central Washington. The other three Ochrogramma species are limited to small areas of Oregon and northern California (Gardner and Shelley 1989, Fig. 220). The meaning of the name Ochrogramma was not discussed by Gardner and Shelley (1989). Both parts of the name come from Greek; ochros a prefix meaning “ yellow ” and gramma, referring to a “ line, writing or drawing ” (the root of English words such as grammar, graphics, telegraph, etc.). The combination likely refers to the dorsal yellow stripe.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFE18B583580FA09FED1F872.taxon	description	Descriptive notes. Gardner and Shelley did not report any records of Ochrogramma species north of the Columbia River, but given the wide distribution of O. bentona in western Oregon — the northern three fourths of the coastal region to just south of the river — our new records are not surprising. These records extend the distribution of the species north to the Puget Sound region. We found little variation in the gonopods of the specimens we examined, but Gardner and Shelley (1989) illustrated a wide degree of variation in the female vulvae or cyphopods (see their Fig. 38 – 41), raising the possibility of more than one species being present; alternatively, the female genitalia could be quite variable within the species. Examination of the DNA of freshly collected material would answer this question. As with males of V. irritans, O. bentona has the typical suite of caseyid modifications of the anterior legs and legpair 10 (Fig. 15 – 18). Legpair one shows reduced telopodites with longer, flattened setae on the postfemora and tibiae. Legpair two coxae bear long, thin, acutely pointed gonapophyses that are longer than the legpair two telopodites (ga, Fig. 15); the openings of the vasa deferentia are large and just posterior to the bases of the gonapophyses (vd, Fig. 15). In many preserved specimens the gonapophyses are carried in front of the male’s “ face, ” but we do not know if this is the case in life. Legpair three coxae are extended into the usual broad lobes that do not exceed the somewhat reduced telopodites (cx 3, cl, Fig. 16). The setae on the tips of the lobes arise from deep sockets, have somewhat swollen bases and are variously curled and appressed to the surface of the lobe (Fig. 17). The coxae of legpair seven (cx 7, Fig. 18) each have a low, flattened mediodistal lobe. Ochrogramma bentona gonopods (Fig. 19 – 24) are considerably simpler than those of V. irritans, principally because the number of arborescent processes is greatly reduced; these are limited to a small area near the apex of the anterior angiocoxite as seen in anterior view (aac, Fig. 19). There appears to be a median, quadrate sternal sclerite linking more massive lateral lobes on either side, with a partial separation by a membranous region. In lateral view (Fig. 21), the anterior angiocoxite (aac) is apically sigmoidally twisted and bears on its mesoposterior surface a large, complexly branched process (Fig. 23). Arising near the base of the angiocoxite is a similar, more sparsely branched structure that is probably the flagellocoxite (f, Fig. 21). The much reduced telopodite (t, Fig. 20, 21) articulates at the lateral base of the anterior angiocoxite and is divided at its tip into several slender branches. The posterior angiocoxite (pac, Fig. 20 – 22) is slender, rodlike, and as with the telopodite is finely divided apically. The colpocoxite (cc, Fig. 20 – 22) is somewhat more robustly sclerotized than usual and in keeping with its hypothesized origin from the coxal gland of the eighth legpair is seen in posterior view (Fig. 20) to arise from a socket or pore. Interestingly, the arborescent processes of the anterior angiocoxite have a stereotypical form (Fig. 24), each with three divisions. The most basal is a low, rounded knob with four or five “ warts, ” the next distal division a larger version of the most basal, and the most distal division a long, arched stem with many branches, but the branches do not end in the cuplike tips of those of V. irritans. The ninth legs (Fig. 25, 26) are typical of caseyids, with coxae (cx 9, Fig. 25) bearing mesal processes (Fig. 26) and single-articled, button-like telopodites (t 9, Fig. 25). The coxae seem to have a more lightly sclerotized area on their mesal surface just proximal to the processes, but no pores could be detected. Legpair 10 (Fig. 27) is very much as in V. irritans and this form, with a double process from the trochanter (trp, Fig. 27), which is partially fused to the coxa (cx 10, Fig. 27), might be a subfamilial character. The opening of the coxal gland (cg, Fig. 27) is very large and occupies much of the mesal surface of the coxa. The female genitalia are illustrated in posterior view in Fig. 31. The outer (ov) and inner (iv) valves appear to be fused. Ochrogramma bentona was recorded by Gardner and Shelley (1989) from the following Oregon counties: Benton, Coos, Lincoln, Polk and Washington. In addition to the new records from Washington state, we add the Oregon counties of Clackamas, Columbia and Linn. New records OREGON: Benton Co.: Missouri Bend Recreation Area, N 44 ° 20.68 ′, W 123 ° 43.17 ′, 229 ′ asl, 29 September 2004, W. Leonard, m; 1 mi on Botkin Road from SR 34, N 44 ° 27.826 ′ W 123 ° 27.701 ′, 25 February 2005, W. Leonard, C. Richart, m, f; Clemens Park, Seely Creek Road, 0.3 mi from SR 34, North Fork Alsea River, N 44 ° 24.55 ′, W 123 ° 34.07 ′, 400 ′ asl, 4 December 2005, W. Leonard, C. Richart, mm, f. Clackamas Co.: Mount Hood National Forest, Ripplebrook Campground, N 45 ° 04.801 ′, W 132 ° 02.542 ′, 1420 ′ asl, 26 October 2003, W. Leonard, m. Columbia Co.: Keystone Creek, Clatskanie Valley, N 40 ° 04.933 ′, W 123 ° 09.411 ′, 132 asl, 8 February 2004, W. Leonard, C. Richart, m, ff. Lincoln Co.: Along SR 34 between Tidewater and Waldport, 19 November 1946, J. C. Chamberlin, m, f (FSCA). Linn Co.: Willamette National Forest, Sweet Home Road, 1 June, 3 August, 12, 27 September, 1 October 2001, J. Rykken, mm, ff. WASHINGTON: Cowlitz Co.: SR 504, 2.1 mi E of Toutle, N 46 ° 20.90 ′, W 122 ° 42399 ′, 500 ′ asl, 1 March 2004, W. Leonard, C. Richart, mm, ff. Grays Harbor Co.: 3 mi N on SR 12 on Middle Satsop Road, N 47 ° 01.992 ′, W 123 ° 31.553 ′, 26 November 2004, W. Leonard, m; N 47 ° 18.879 ′, W 123 ° 33.382 ′, 830 ′ asl, 13 October 2003, W. Leonard, m. King Co.: Exit 18 on I- 90, near Olallie State Park, N 47 ° 26.651 ′, W 121 ° 40.081 ′, 23 February 2004, W. Leonard, C. Richart, m. Lewis Co.: 8 mi S of Randall, 6 December 2003, W. Leonard, C. Richart, m, ff; 8.5 mi S of Randall on FS 25, Gifford Pinchot National Forest, W 46 ° 26.489 ′, W 121 ° 59.890 ′, 21 December 2003, W. Leonard, mm, ff; Iron Creek Campground, 9.6 mi S of Randall on FS 25, Gifford Pinchot National Forest, 950 ′ asl, 21 December 2003, W. Leonard, m, ff, N 46 ° 26.456 ′, W 121 ° 59.796 ′, 1050 ′ asl, 6 December 2003, W. Leonard, C. Richart, mm, ff; 604 Roswell Road, Centralia, N 46 ° 41.339 ′, W 122 ° 56.664, 240 ′ asl, 25 January 2004, C. Richart, W. Leonard, m; SR 508 at Bremer, N 46 ° 15.30 ′, W 122 ° 25.55 ′, 750 ′ asl, 6 December 2003, Q. Leonard, C. Richart, m, f; seep beside road along Cowlitz River east of Riffle Lake, N 46 ° 27.727, W 122 ° 09.763 ′, 650 ′ asl, 9 April, 2004, W. Leonard, C. Richart, m. Pacific Co.: Long Beach, 30 th Street, 0.2 from US 101, N 46 ° 19.436 ′, W 124 ° 03.591 ′, 50 ′ asl, 15 January 2006, W. Leonard, C. Richart, m. Pierce Co.: Sequakitchew Creek trailhead, N 47.107 °, W 122.645 °, 200 – 220 ′ asl, 28 January 2012, R. Crawford, m, f (BMUW). Thurston Co.: McAllister Springs, N 47 ° 02.946, W 122 ° 43.678, 22 February 2004, W. Leonard, m, f N 47 ° 02.837, W 122 ° 43.722, 60 - 120 ′ asl, 7 Feb 2004, W. Leonard, C. Richart, m, ff; McAllister Creek, S of Steelacoom Road, N 47 ° 03.42 ′, W 122 ° 42.79 ′, 100 ′ asl, 11 December 2004, W. Leonard, mm, ff; Hospital Creek, 3 mi S, 3 mi E of Vail, 12 October 2003, W. Leonard, m, at confluence with Skookumchuck River, N 46 ° 46.396 ′, W 122 ° 35.133 ′, 15 December 2003, W. Leonard, K. McAllister, m, ff; Skookumchuck River, 5 mi S, 3 mi E of Vail, 2 October 2003, W. Leonard, mm; Rainier Road, 3.6 mi NW of Military Road, Fort Lewis Military Reservation, 29 November 2003, W. Leonard, mm, ff; Priest Point Park, N 47 ° 04.47, W 122 ° 53.85 ′, 11 January 2004, W. Leonard, m.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFF88B473580FF45FD6BFC60.taxon	description	Descriptive notes. This record (see below) of O. haigi represents a southern extension of the species’ range. It was previously known from Curry Co., Oregon and Del Norte Co., California. Ochrogramma haigi males have the typical modifications in this genus of the mandible (mf, Fig. 32) and pregonopodal legs. The first leg telopodites (t 1, Fig 32) are somewhat reduced, as are those of the second legs (t 2, Fig. 32) and long gonapophyses (g, Fig. 32) curve forward from the second coxae, with prominent vas deferens openings at their bases (vd, Fig. 33). At least in this preserved specimen the gonapophyses are carried in front of the male’s “ face. ” Third leg coxae (cx 3, Fig. 34) are elongate with prominent terminal lobes (cl, Fig. 34). The recumbent setae of the third leg coxal lobes (Fig. 35) have bulbous bases and are striated; the filament of the seta seems to arise beneath the bulbous base. While the arborescent processes of the anterior angiocoxites are present in O. haigi, they are less extensive than in O. bentona as well as less regular in form (Fig. 36). The gonopods are much more simple and apparently lack both the telopodite and the posterior angiocoxite; the flagellocoxites (f, Fig. 36) are tightly appressed to the posterior side of the anterior angiocoxite (aac, Fig. 36). The ninth legpair (Fig. 37) has bulbous coxae (c 9, Fig. 37) with a bifid coxal process (cp, Fig. 37) and a small pore (p, Fig. 37). New record CALIFORNIA: Humboldt Co.: Prairie Creek Redwoods State Park, N 41 ° 22.379 ′, W 124 ° 00.833 ′, 252 ′ asl, 21 December 2006, C. Richart, A. Fusek, m.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFF88B423580FC60FA5FFAF9.taxon	description	(Fig. 38 – 47) Types. Male holotype (VMNH 11244), five male paratypes (VMNH 11245.1 – 5) and 11 female paratypes (VMNH 11246.1 – 11) from Spring Creek Hatchery on State Route 14, 45 ° 43.680 ′, − 121 ° 32.672 ′, 50 ′ asl, Skamania Co., Washington, collected 30 November 2003 by W. Leonard, deposited in VMNH.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFF88B423580FC60FA5FFAF9.taxon	etymology	Etymology. This species has been collected only in Skamania Co., Washington, and the species epithet is a noun in apposition after the name of the county.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFF88B423580FC60FA5FFAF9.taxon	diagnosis	Diagnosis. Ochrogramma skamania, new species, is distinct from the other species of the genus in that the anterior angiocoxite (aac, Fig. 40) of the gonopod is not twisted, while arborescent processes are present on the anterior surface (Fig. 41, 45); the ninth leg coxal processes are not divided as in the other species but instead are entire and squared off distally (cp, Fig. 46).	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
C2099450FFF88B423580FC60FA5FFAF9.taxon	description	Description. Male paratype from Spring Creek Hatchery. Length, ca. 10 mm, width 0.85 mm. Eyepatch of 17 – 19 ommatidia. Mandibular stipes with strongly developed distal lobe. Thirty rings. Rings cylindrical, smooth dorsally, laterally with 10 - 12 prominent horizontal striations. Segmental setae acute, short. Color purplish to brownish gray, with yellow-tan spots around sockets of segmental setae and dorsal yellow-tan stripe more prominent posteriorly. Sides of rings yellowish with striations darker purplish gray. First legpair (Fig. 38) short, arising from a coxosternum (cxs 1, Fig. 38), postfemora, tibiae and tarsi with long, flattened, spirally curved setae. Second legpair (Fig. 39) with coxal gonapophyses (ga, Fig. 39) longer than telopodites (pf 2 et seq., Fig. 39). Third legpair (Fig. 39) with coxae (cx 3, Fig. 39) not fused with sternum (s 3, Fig. 39), elongated into rounded lobes (cl, Fig. 39), distally with specialized setae as in O. haigi (Fig. 35); telopodites (t 3, Fig. 39) articulating about two-thirds distal on coxae. Seventh legpair with short coxal lobes. Gonopods (Fig. 40 – 45, 49) with coxae and sternum (s, Fig. 40, 41) clearly separate, coxae (c, Fig. 40, 41) rounded, cupulate. Anterior angiocoxites (aac, Fig. 40 – 45) slender, curved, slightly expanded and rugose at tips, with arborescent processes on anterior distal surface. Flagellocoxites (f, Fig. 42, 43) three, long, branched, almost feathery, appressed to posterior surface of anterior angiocoxite. Telopodites absent. Posterior angiocoxite (pac, Fig. 42 – 44) rodlike, apically divided into several fine filaments. Colpocoxites (cc, Fig. 40 – 43) weakly sclerotized, arising from coxal sockets, directed mesally. Ninth legpair (Fig. 46) with coxae (cx 9, Fig. 46) free from sternum, each bearing long, unbranched mesal process (cp, Fig. 46), curved and truncate at tip; prominent pore (p, Fig. 46) at anterior base of each coxal process. Tenth legpair coxae (cx 10, Fig. 48) much enlarged, with large, anterior gland openings (cxg, Fig. 48); trochanters (tr 10, Fig. 48) partly fused to coxae, distally with bifid, blunt process bearing many small teeth (trp, Fig. 48). Female paratype from Spring Creek Hatchery. Length, ca. 15 mm, width 1.2 mm. Nonsexual characters as in male, but dorsal yellow stripe more prominent, extending further anterior. Female genitalia (Fig. 47) with widely separated outer (ov) and inner (iv) lobes, outer lobes shorter, both lobes with strong distal processes. Receptacles not observed, postgenital plate absent. Records WASHINGTON: Skamania Co.: Cape Horn, 45 ° 34.29 ′, − 122 ° 11.86 ′, 400 ′ asl, 19 November 2004, W. Leonard, m, ff (VMNH); Cook-Underwood Road, 1.4 mi northeast of Cook, 45 ° 434 ′, − 121 ° 38.859 ′, 450 ′ asl, 30 November 2003, W. Leonard, f (VMNH). Note. This is the first new species of Ochrogramma from north of the Columbia River. Spermatophores and Spermatozoa Millipede spermatocytes and spermatozoa were reviewed by Minelli and Michalik (2015), among others. Millipede spermatozoa are highly variable in form from order to order, but all are evidently nonmotile. According to Minelli and Michalik (2015), no anatomical or ultrastructural studies have been carried out on any nematophoran taxa, so our observations here, however sketchy, are the first data on spermatozoa of this large taxon which includes the orders Callipodida and Chordeumatida. Our SEM pictures only show spermatozoa embedded in a matrix, evidently comprising a spermatophore (Fig. 28). They appear to be subglobular or slightly flattened (most closely resembling the spermatozoa of the order Julida, based on the illustrations in Bacetti et al. [1979]) and 14 – 14.5 µm in diameter. There is no indication of any organ of motility. Minelli and Michalik (2015) make no reference to spermatophores, nor do they in a discussion of mating behavior immediately following. This is curious, as the paper by Mathews and Bultman (1993) clearly discusses a spermatophore in the mating of the parajulid (Julida) Aniulus bollmani Causey, 1952 (= Aniulus garius (Chamberlin, 1912). This is not the place for a review of either the evidence for, or speculation about, spermatophores in chordeumatidan millipedes, aside from positing the idea that sperm from the vas deferens, opening on the second coxae, are first transferred to the large coxal glands on the tenth legs where they are formed into spermatophores by a secretion. The spermatophores would then be picked up by the gonopods and by them inserted into the female vulvae during mating. Supposed spermatophores have already been illustrated in, for example, Trichopetalum dux (Chamberlin, 1940) on both the gonopods and attached to the female vulvae (Shear 2003). In that case the spermatophores seemed to have a characteristic shape. In the course of our SEM studies of Ochrogramma species, we found what we consider to be amorphous spermatophores or spermatophore material located in the tenth legpair coxal glands of males (sp, Fig. 48) on the gonopods (sp, Fig. 30, 49) and attached to the vulval valves of females (sp, Fig. 31, 47). This is strong evidence that the theoretical sequence described in the paragraph above accurately describes what happens in the mating of chordeumatid species. The secretion forming the spermatophores is extremely sticky, or may have been hardened by preservation in alcohol. As can be seen from the illustrations, it could not be removed by ultrasonic cleaning.	en	Marek, William A. Shear Paul E. (2024): Additions to the millipede family Caseyidae Verhoeff, 1909. III. Proposal of the new subfamily Ochrogrammatinae, new records of Vasingtona irritans (Chamberlin, 1910), established species of Ochrogramma Gardener and Shelley, 1989 a new species of Ochrogramma from Washington state, USA. Insecta Mundi 2024 (90): 1-21, DOI: 10.5281/zenodo.14662376
