Guestphalinus elephantinus Fend & Rodriguez & Achurra & Erséus 2017, sp. nov.

Guestphalinus elephantinus Fend & Rodriguez sp. nov. urn:lsid:zoobank.org:act: A30D385C-06C9-438E-AAFC-1F145905B74F Figs 10–12 “ Guestphalinus sp. nov.” – Zhou et al. 2010: 381, figs 1–6 (sequenced, but not morphologically described; voucher SMNH 105628). Etymology Based on the fanciful and homoplasious similarity to an elephant (proboscis and anteriorly-directed “tusks”). Material examined Holotype UNITED STATES OF AMERICA: a dissected specimen, slide-mounted in Canada balsam, Washington, Jefferson County, Shale Creek at Clearwater Creek Road, 4 Jun. 2003, S. Fend leg. (USNM 1422286). Paratypes (all from the Clearwater River drainage, on Olympic Peninsula, Washington) UNITED STATES OF AMERICA: 1 dissected (post-reproductive), Jefferson County, Clearwater River at Upper Clearwater Camp, 27 Apr. 2004 (USNM 1422287); 1 dissected (post-reproductive), same data as preceding (CASIZ 220932); 1 dissected (post-reproductive), Hurst Creek near Clearwater River, 25 Apr. 2004 CASIZ 220933); 1 whole mount, immature but with gonads (DNA voucher), from type locality, 4 Jun. 2003 (CASIZ 220934); 1 dissected (post-reproductive), Clearwater River at Upper Clearwater Camp, 27 Apr. 2004 (MNCN 16.03/3098); 2 partially-mature whole mounts on 1 slide, Shale Creek at Clearwater Creek Road, 4 Jun. 2003 (MNCN 16.03/3099). Additional material (typical specimens) UNITED STATES OF AMERICA: WASHINGTON, Jefferson County (all from the Clearwater River drainage, on Olympic Peninsula): 2 dissected (mature, reproductive organs slightly resorbed), Bull Creek near Clearwater River, 2 Apr. 2016; 2 whole mounts (immature, with small gonads), Clearwater River at Upper Clearwater Camp, 4 Jun. 2003; 4 dissected, (immature or post-reproductive), same locality as preceding, 27 Apr. 2004; 1 dissected and 16 whole mounts, several in alcohol (all immature or partially mature), Clearwater River at Copper Mine Bottom Camp, 25 Apr. 2004; 8 whole mounts (immature or partially mature), Shale Creek at Clearwater Creek Road, 4 Jun. 2003; 4 whole mounts (immature), same locality as preceding, 14 Apr. 2010; 5 whole mounts (immature), Hurst Creek, 25 Apr. 2004. Other sites UNITED STATES OF AMERICA: WASHINGTON: 1 dissected (slightly post-reproductive), 2 whole mounts (post-reproductive), Grays Harbor County, Middle Fork Satsop River, 27 Apr. 2004, S. Fend leg.; 2 whole mounts (1 nearly mature), Columbia County, Tucannon River below Turner Road, 29 Jul. 2011, Uttam Rai leg. – OREGON: 1 sagittal section, 1 transverse section, 3 dissected, 6 whole mounts (dissected and sectioned nearly mature, others with gonads),Yamhill County, spring at Peavine Ridge near McMinnville, 5 Dec. 1999, S. Fend leg.; 1 whole mount (immature, with gonads), Curry County, Mule Creek at upstream bridge, near Rogue River, 7 Jun. 2003, S. Fend leg.; 1 dissected (immature, with gonads), Curry County, Euchre Creek near mouth, 20 May 2013, S. Fend leg. – CALIFORNIA: 1 whole mount (immature), Santa Clara County, Alamitos Creek above Almaden Reservoir, 1 May 1997, S. Fend leg.; 1 whole mount (immature), Santa Clara County, Guadalupe Creek above Guadalupe Reservoir, below Rincon Creek, 3 Apr. 2001, S. Fend leg.; 1 dissected (mature) and 2 whole mounts (partially mature), same locality as preceding, 25 Mar. 2007, S. Fend leg.; 1 whole mount (immature), same locality as preceding, 10 Feb. 2008, S. Fend leg.; 1 whole mount (immature), same locality as preceding, 5 Dec. 2009, S. Fend leg.; 1 dissected (post-mature), same locality as preceding, 20 Jun. 2011, S. Fend leg. Molecular data COI, 28S and 16S sequences are based on specimens from Shale Creek and Clearwater River, Washington (details in Table 1). An additional ITS sequence (GenBank acc. no. GU592364) corresponds to the Clearwater River voucher SMNH 105628. A topotypic voucher is included in the type series (see above). Both COI sequences for G. elephantinus sp. nov. are identical. Description Specimens from the Clearwater River drainage, Olympic Peninsula, Washington Description of somatic characters is based on mature or partially-mature (with gonads) worms from Clearwater River, Hurst Creek, Shale Creek and Bull Creek. Reproductive organs are described from the only fully-mature specimen (the holotype), from Shale Creek, and supplemented with 9 postreproductive specimens with partially-resorbed organs from nearby sites Body form and segmentation as described for G. exilis sp. nov. (see above), measurements given in Table 3. Length of proboscis 0.9–2.6 mm (Figs 11A, 12A), clitellum from VIII to XII in the mature worm. Chaetae in II much larger and thicker than those in posterior segments (Table 3), bifid, with a short, thick lower tooth and a larger, rounded and laterally compressed upper tooth (Figs10 A–C, 12C), directed anteriorly, with proximal ends protruding into posterior segments (cf. Figs 11 C–D, 12B). Dorsal chaetae in II about 30–40% longer than those in ventral bundles (Table 3); within each bundle in II, the lateral (i.e., outer) chaeta is slightly longer, with a more distal nodulus (0.15 from tip, vs 0.24 in the median (inner) chaeta), and usually has a distal thickening that appears as a “secondary nodulus” between nodulus and tip (Fig. 12 C–D). Chaetae in III simple-pointed or slightly bifid, and oriented more perpendicularly to body, but still facing anteriad (Figs 10C, 12B); much shorter than chaetae in II and slightly shorter than those in other segments (Table 3). Chaetae in IV and posterior segments usually oriented posteriorly, simple-pointed, sigmoid (Fig. 12H). From III to about XX, nodulus about 0.3–0.4 from the tip, but the median (inner) chaeta in each bundle is slightly longer. In posterior segments, chaetae of similar length, but slightly thicker, and nodulus 0.25–0.35 from the tip; simple-pointed or with a small notch or dorsal keel in tail segments. Ventral chaetae absent in IX in mature or post-mature worms. Epidermis in anterior segments 15–20 µm thick; in clitellum to 48 µm in the holotype; narrower, densely packed cells up to 30 µm on ventral side of IX (in area surrounding genital pores) in the holotype and in the post-reproductive worms; posteriorly 10–12 μm. Body-wall musculature, brain, pharynx, pharyngeal glands, nephridia, blood vessels and chloragogen as described above for G. exilis sp. nov., except that pharyngeal glands in G. elephantinus sp. nov. extend to VII in all specimens observed. Male and spermathecal pores paired in IX. Male pores just anterior to the posterior intersegmental groove (Fig. 11A), slightly inside the line of ventral chaetae; spermathecal pores aligned with and in front of male pores, between or slightly posterior to chaetal bundles. Female pores paired, on chaetal line at 10/11; female funnel up to 130 µm high (Fig. 12Q). A variable number of copulatory glands with internal structure as described for G. exilis sp. nov.; with round secretory openings located close to spermathecal and male pores. The mature worm (holotype) has 3 pairs of petiolate copulatory glands in IX, 100–140 µm high, lateral to, in front of, and behind genital pores (Fig. 11E), although not all of these were visible externally (Fig. 11A). Copulatory glands smaller and indistinct or absent in the postreproductive worms (male duct and spermathecae expanded; large but nearly empty sperm and egg sacs extending to XV), but faint external secretory openings visible in one specimen. Spermathecae up to 1700 µm long in the holotype; ectal ¼ is duct-like, nearly tubular (about 90 µm in diameter); ental ampulla-like portion irregular, up to about 100 µm in diameter, folded within X or extending into XI. Ectal part with irregularly columnar cells up to 35 µm high, epithelium gradually becoming thinner in ental ampulla. Spermathecae end in a short, narrow constriction surrounded by a ring of muscle fibers (Fig. 11E, cf. Fig. 12L) which opens in a shallow epidermal infolding, about 40 μm deep. Spermathecae of a paratype similar (Fig. 11G), 1200–2600 µm long, with loose sperm; ampulla thin-walled, ental diameter up to 230 µm, extending into X or as far as XII. Anterior pair of male funnels on 8/9; posterior male funnels on 9/10, directed back into X; male funnels of post-reproductive worms indistinct, without or with small amount of sperm. Anterior vas deferens extends into VIII (Fig. 11E), forming a compact, convoluted mass, then penetrates 8/9, running along ventral body wall to near the male pore, then follows the atrium before joining the atrial wall and entering the lumen near the apex (junction not clear on holotype). Length of anterior vas deferens up to about 1500 μm, width of both anterior and posterior vasa deferentia 30–36 μm in the holotype. Posterior vas deferens forms a compact, convoluted mass in posterior part of IX, then follows the atrium externally before joining it near the ental end. Atria of holotype extend back into X; ectal duct (length 160 μm, width near midpoint 30–40 µm) has a thick, columnar epithelium and thin muscle coat; ectal part of duct expanded to 50 µm, with epithelial cells directed outward. Entally, there is an abrupt transition to the sacciform atrial ampulla; ampulla length up to 550 μm, width up to 130 μm, with thin-walled epithelium and a wide lumen (Fig. 11E). Small clusters of prostatic cells, 20–40 µm high, very sparse. Atria tubular, with ampulla not expanded in a paratype (apparently at a more advanced stage of maturity, after copulation, Fig. 11G); length 400–800 µm, diameter 60–80 µm; prostates very small or absent; atrial duct and male pore as in the holotype. Specimens from other localities Satsop River, western Washington: Modified chaetae in both II and III; chaetae in II very long (Table 3); lateral (outer) chaeta in the largest specimen with secondary nodulus, as in topotypic specimens (Fig. 12D). Chaetae in III smaller, but bifid and larger than chaetae in the next segments, and directed anteriad (Fig. 11D). Tucannon River, eastern Washington: Typically modified chaetae

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