identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
154FD01D4B51FFD4FF6E92AFBBE3CBA8.text	154FD01D4B51FFD4FF6E92AFBBE3CBA8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus saelonae	<div><p>Rhyacodrilus saelonae sp. n.</p><p>(Figs 1–3)</p><p>Holotype. USNM 1202047, whole-mounted worm stained in hematoxylin and mounted in Canada balsam.</p><p>Paratypes. USNM 1202048-52: 3 mature specimens from the type locality, stained in hematoxylin or borax carmine and whole-mounted (17 March 2008); 1 mature, dissected worm, stained with hematoxylin (7 May 2006); 1 sagittally sectioned (17 March 2008). MNCN 16.03/3079: 3 mature specimens from the type locality, stained in borax carmine and whole-mounted (17 March 2008).</p><p>Type locality. Nacimiento River, above Fort Hunter Liggett, at upstream bridge, Nacimiento-Fergusson Road, Monterey Co., California, USA. N36.0132°, W121.4201° (17 March 2008).</p><p>Other material. From the type locality: 1 whole mount (22 April 2006), 8 whole mounts and 1 dissected (7 May 2006), 2 sagittally sectioned and 13 whole mounts (17 March 2008). Nacimiento River below Negro Fork, N35.9969° W121.3797°, 6 whole mounts (3 April 2008).</p><p>Etymology. The species is dedicated to Saelon Renkes, who has contributed to the discovery of many Nearctic oligochaete species.</p><p>Description. (Measurements are based on individuals with sperm in spermathecae, with or without welldeveloped eggs). Number of segments 65–96 (based on 5 complete specimens). Diameter of the body in segment VIII (whole-mounted specimens) 217–362 µm. Elongated, round prostomium (197–240 µm long) (Figs 1 A, 2A). Secondary annulation clearly marked in II; faint but visible in other preclitellar segments of most specimens. Epidermis not densely glandular (8–12 µm high), small glandular cells unordered. Clitellum from the level of chaetae in the spermathecal segment to the end of the post-atrial segment, thickest dorsally and laterally, weak near male and spermathecal pores; clitellar epidermis up to 47 µm high (in most specimens about 20–30 μm), formed by large glandular cells in the dorsal and lateral part, extending ventrally in the post-atrial segment; ventral epidermis thickened but not glandular in the spermathecal segment, and thin in the area between the male pores (Figs 1 A, B, 2C).</p><p>Dorsal bundles composed of smooth hair-like and pectinate chaetae. Hair chaetae (1)2–3 per bundle anteriorly, and (0)1–2(3) in postclitellar segments, much longer than body diameter (408–800 µm) in the anterior segments, and longer or about the same as body diameter in posterior segments. Usually 3–4 pectinate chaetae in anterior bundles (rarely 2), and (1) 2–3 in postclitellar segments; 67–111 µm long (much shorter in II, down to 41µm). Pectinates in anterior segments with lyre-shaped lateral teeth (7–13 µm long) and 2–4 shorter (4–8 µm) intermediate teeth; pectinates in posterior segments with shorter lateral teeth (6–8 µm long) (Fig. 2 D, E). The last segments of complete specimens with only 1 pectinate chaeta (48 µm long) per bundle, with very short teeth, and without hair chaetae. Ventral chaetae bifid; 5–9 per bundle (rarely 3 in segment II) in anterior segments, 4–7 in postclitellar segments; distal tooth 1.6–1.8 times longer than proximal in anterior segments, becoming shorter (about same length) and thinner posteriorly (Fig. 2 F, G). In the last segments down to 1–3 ventral chaetae per bundle, with distal tooth shorter and thinner than proximal.</p><p>One pair spermathecal pores; simple, oval openings, up to midway between ventral chaetal line and lateral line, near anterior septum of the testicular segment. Modified spermathecal chaetae behind spermathecal pores, (3)4–5 per bundle (commonly, one of them developing), in line with ventral chaetae, sometimes on a low mound with cellular extensions from the ventral gland penetrating the epidermis behind chaetae. Spermathecal chaetae 116–158 µm long and 4 µm shaft diameter, with very long, sharply-pointed, grooved distal tooth (14–19 µm long); the short, proximal tooth appears as a wide prominence at the base (Figs 2 H, I, L). Nodulus 0.3–0.4 from the distal end. Two ridges of raised epidermis, U-shaped to nearly circular, open laterally (diameter 80–100 μm, 20–35 μm high at midline) and conjoined at ventral midline between the spermathecal pores, forming an "anchorage bridge" (sensu Cuadrado and Martínez-Ansemil, 2001). A conspicuous, transverse epidermal fold (about 80–120 μm wide, up to 30–50 μm deep) opens at septum behind (and in line with) spermathecal chaetae.</p><p>One pair simple male pores in the ovarian segment, on (or slightly median to) the ventral chaetal line, at about the posterior 1/4 of the segment. Modified penial chaetae slightly median to the male pore, in the atrial segment, single-pointed, 120–161 µm long, 3–4 per bundle, weak nodulus at 0.2 from distal end (Fig. 2 M). Penial chaetae all of the same size or the middle ones slightly longer; arranged roughly fanwise, with tips close together and directed medially; usually on low tubercles (about 25 µm high); the tubercles and chaetae surrounded by a ring of muscles (Fig. 1 F). A complex of dorso-ventral muscular strands is also associated with the penial chaetae, attaching to the tubercle near the tips of the chaetae; bases of chaetae have the usual chaetal musculature attaching them to the ventral body wall. One pair female pores at posterior intersegment of ovarian segment, midway between ventrolateral chaetal line and lateral line. Reproductive organs are often shifted forward by 1 or 2 segments relative to typical Rhyacodrilus, but they retain their relative positions: spermathecal pores in VIII, IX, or X correspond to male pores in IX, X, or XI, respectively, and the clitellum begins in the spermathecal segment.</p><p>Nucleated coelomocytes with granulated cytoplasm (diameter 12–18 µm) (Fig. 2 B). Pharyngeal glands welldeveloped in IV–V, usually extending into VI. Nephridia paired, single, or absent in the postclitellar segments, sometimes in one or more preclitellar segments; with short, narrow anteseptal funnel, an elongate (sometimes inflated), granular postseptale (Fig. 2 J), and long efferent duct which may extend posteriorly into the next segment; the duct terminates at an inconspicuous nephropore just in front of the ventral chaetae. One pair ventral glands behind the ventral bundles of chaetae, from segment (IV)V to the spermathecal segment, and from the post-atrial segment as far back as XX; the largest in the spermathecal segment (up to 140 μm wide), well-developed in prespermathecal and post-atrial segments (50–90 μm wide), then progressively smaller. These glands project freely into the coelomic cavity and the cell necks extend through the muscular layers and body wall to the body surface forming an area glareosa (sensu Schmelz 2003: 49f) visible in vivo (Figs 1 E, 2K, 3A–C ag). Sperm sac may extend anteriorly to segments VIII and IX, and backwards to segment XIII. Egg sac may extend back to segment XIV.</p><p>One pair testes in the spermathecal segment (typically IX) and one pair ovaries in the atrial segment (typically X). Spermathecae composed of a short, conical duct, usually shorter than wide (34–60 µm long, 48–62 µm maximum diameter), with a thick wall of columnar epithelium, and an oval ampulla which occupies most of the length of the segment (120–282 µm maximum diameter in mated worms), with thin epithelium (6–12 µm high). Sperm free within lumen of spermathecal ampulla, but anterior ends of spermatozoids may be oriented towards one end of the ampulla. One ampulla may penetrate into an adjacent segment. Several specimens show in lateral view a dorsal fold in the ampulla (Fig. 3 D).</p><p>Male duct opens into a conical sperm funnel; vas deferens highly convoluted, ciliated, and of nearly uniform width (diameter 24–37 µm), length 530–750 μm, running along the ventral side of the segment and narrowing as it joins the atrium subapically, or in some specimens near the middle of the ampulla. Atrial ampulla ovate (95–175 µm long, 54–96 μm wide), completely covered by a thick (30–60 μm) layer of prostate cells, at least some of which are in densely packed clusters of 10 or more cells. Ampullar epithelium appears irregular and vacuolated, to 25 μm thick, sometimes nearly filling the atrial lumen (Fig. 3 F–H). Atrial duct 100–194 µm long, 43–52 µm wide), slightly narrowed entally, with a uniform, columnar epithelium, narrow lumen, and thin outer muscle layer; a dense array of muscular strands runs from ventral body wall to the outer muscle layer along the entire length of the duct (Fig. 1 C, D). Female funnels long, in some specimens more than half the diameter of the body (Fig. 3 F, G).</p><p>Anomalies. Positions of reproductive organs vary considerably in this species. More than half of the specimens had the genitalia shifted one or two segments forward from the usual positions in the Rhyacodrilinae . Of 36 mature worms examined, 12 had the spermathecae in X, 16 in IX, and 8 in VIII; in all cases the relative position of other reproductive organs to the spermathecae was preserved, and both male ducts and spermathecae were functional (with sperm). Some species of the genus Rhyacodrilus have been reported to have architomy followed by regeneration, which may be associated with a shift in the position of the reproductive segments (e.g. Snimschikova 1985), although we have not seen any evidence of developing anterior segments in this species. Two specimens had enlarged ventral chaetae in the post-atrial segment with upper teeth much longer than lower, similar to the spermathecal chaetae (but smaller); in this same segment there was also a small gonad and a slightly enlarged ventral gland behind the chaetal bundle. One specimen had an extra spermatheca on one side of the body, at the atrial segment (X), but only the spermatheca in the preatrial segment was filled with sperm. Another specimen had a single spermatheca (with sperm) in VII, a rudimentary spermatheca in VI, one pair spermathecae with a single functional male duct in VIII, and functional male ducts paired in IX.</p><p>Distribution and habitat. Known only from type locality, a largely undisturbed section of the Nacimiento River, in the Santa Lucia Mountains (Los Padres National Forest). The two reaches with R. saelonae were in a small stream with permanent surface flow, near the transition from high to low gradient sections. Sediment was a mixture of colluvial and alluvial particles, sizes ranging from cobbles to coarse gravels, overlaying coarse sand. The sampling area is a salmonid habitat, with clear, cool water, a well-preserved riparian habitat and little aquatic vegetation.</p><p>Remarks. Based on the relative position of the reproductive organs (spermatheca in the most anterior part of the testicular segment, plesioporous male duct and male pores opening in the ovarian segment), the prostate diffuse or with cells in clusters broadly attached to the atrial surface (Erséus 1984), and the abundance of large nucleated and granulated coelomocytes, the new species is member of the subfamily Rhyacodrilinae . Although this subfamily requires revision due to its paraphyletic nature (Erséus et al. 2008, Martin et al. 2010), the new species has all the main diagnostic characteristics of the genus Rhyacodrilus: moderately long vasa deferentia entering atria subapically, well-developed prostate glands covering the atrial ampulla, coelomocytes present, and modified penial chaetae (Brinkhurst &amp; Jamieson 1971).</p><p>The group of Rhyacodrilus species having both spermathecal and penial chaetae is summarized in Table 1. Among these species, only two were previously known that share both hair and pectinate chaetae in dorsal bundles and a well-developed layer of prostate cells: R. svetlovi Sokolskaya, 1976 (Chukchi Peninsula, Siberia) and R. tauricus Dembitsky, 1975 (Crimea, Ukraine). The former differs from R. saelonae sp. n. in several respects: It lacks hair chaetae in segments posterior to XV, the atria are tubular, the vas deferens is shorter than the atrium, the atrial duct is very short, and the vas deferens joins the atrium apically. The new species is separated from R. tauricus by the shape of the spermathecal and posterior dorsal chaetae as well as by the junction of vas deferens, which is on the posterior face of the atrium in R. tauricus . Modified spermathecal-like chaetae are present in both preatrial and postatrial segments in R. tauricus . The general structure of the male duct in R. saelonae sp. n. (i.e. atrial duct as long as or longer than atrial ampulla, and long vas deferens entering atrium subapically), together with the presence of both penial and spermathecal chaetae, the extraordinarily long distal tooth of the spermathecal chaetae, the lateral position of spermathecal pores, and the presence of ventral glands clearly distinguish this species within the genus Rhyacodrilus .</p><p>The ventral glands in R. saelonae sp. n. are similar in structure to Timm's glands, described in Haplotaxidae by Hrabĕ (1931), and are another diagnostic character for the new species. In our collection, partially mature R. saelonae have small glands in the spermathecal segment which are similar in size to the ventral glands in other anterior segments (based on the examination of 8 specimens at varying stages of reproductive development). The glands in the spermathecal segment are well-developed (larger than those in adjacent segments) in two mature but unmated specimens. This development of the glands in the spermathecal segment during maturation, as well as their position close to the spermathecal chaetae, suggests some role during mating.</p><p>The presence of modified chaetae in the postatrial segment is extremely rare among rhyacodrilines, although they have been reported in Protuberodrilus Giani and Martínez-Ansemil, 1979 and Rhyacodrilus tauricus . In R. saelonae sp. n., this occurred in only two specimens and is interpreted as an anomaly.</p><p>Among Rhyacodrilus species, a lateral position of the spermathecal pores has been reported for R. gasparoi Martínez-Ansemil et al., 1997, a cave species from Italy. That species also has well-developed ventral glands in segment X, associated with spermathecal chaetae. The distinct transverse folds between the spermathecal and atrial segments of the new species correspond to the position of the spermathecal chaetae in the expected inverted copulatory orientation, and thus may have a copulatory function. Other external copulatory structures of R. saelonae appear to be a variation on those described by Cuadrado and Martínez-Ansemil (2001) for Peristodrilus montanus (Hrabĕ, 1962) . The relative positions of the anchorage bridge (here formed by two concave tubercles), the penial chaetae, and male and spermathecal pores are similar. The low tubercles and dorso-ventral muscles associated with penial chaetae may correspond to the "raised ventral elevation" and associated musculature described by Cuadrado and Martínez-Ansemil (2001) for P. montanus . The ring of circular muscles surrounding the penial chaetae in R. saelonae was not mentioned in the latter account, but would have the function of pulling the tips of the chaetae (and the tubercles) together. As the bases of these chaetae are fixed by normal chaetal musculature, the effect would be forceps-like.</p></div>	https://treatment.plazi.org/id/154FD01D4B51FFD4FF6E92AFBBE3CBA8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
154FD01D4B54FFD8FF6E957EBBD7CE3B.text	154FD01D4B54FFD8FF6E957EBBD7CE3B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus quileuticus	<div><p>Rhyacodrilus quileuticus sp. n.</p><p>(Figs 4–5)</p><p>Holotype. USNM 1202053: whole mounted specimen stained in borax carmine and mounted in Canada balsam. Paratypes. USNM 1202054-55: 2 whole mounted specimens, stained in borax carmine, from the type locality (29 April 2004).</p><p>Type locality. South Fork Calawah River, Olympic National Forest, Clallam Co., Washington, USA, N47.9627° W124.2941° (29 April 2004).</p><p>Other material. 4 mature whole-mounted worms (29 April 2004). 1 mature whole mount, stained in hematoxylin, from West Fork Humptulips River, Washington, Grays Harbor Co., approx. N47.48° W123.68° (2 June 2003).</p><p>Etymology. the species has been named after the Quileute People (also called Quilayute) of the Calawah River basin.</p><p>Description. Number of segments: more than 25 (all specimens incomplete). Diameter of the body in segment VIII (whole-mounted specimens) 276–358 µm. Round prostomium (148–217 µm long) (Fig. 4 A). Epidermis 7–12 µm high, and up to 30 µm high at the clitellum, which extends from the level of chaetae in segment X to the end of XII (Fig. 5 A). Clitellum formed by large glandular cells (Fig. 4 C) except for the ventral side around male pores, where there are only small glandular cells. One pair male pores in XI in line of ventral chaetae. One pair spermathecal pores in X, slightly lateral to the line of ventral chaetae, close to intersegment 9/10. One pair female pores opening in intersegment 11/12, about midway between ventral chaetae and the lateral line.</p><p>Hair chaetae (0)1–2 per bundle anteriorly, and 0(1) in postclitellar segments, usually absent posterior to segment XII or XV, and shorter than body diameter (length 110–200 µm). Dorsal pectinate chaetae 2–4 per bundle, nodulus 0.3–0.4 from distal end, 64–73 µm long (56–60 µm in II), distal tooth longer or about equal to proximal, intermediate teeth same length or shorter than lateral teeth (Fig. 4 D, E). Ventral chaetae bifid; anterior ventral chaetae 3–6 per bundle, 75–82 µm long (58–70 µm in II), distal tooth twice length of proximal; 3–5 ventral chaetae in postclitellar segments, 58–76 µm long, with distal tooth thinner, but about the same length as proximal (Fig. 4 F– H). Nodulus at 0.4–0.5 from distal end in anterior ventral chaetae, 0.3–0.5 in posterior ones.</p><p>Modified spermathecal chaetae in segment X, 1–2 per bundle, in line with ventral chaetae, in the posterior third of the segment, well-separated from spermathecal pores (75–97 µm long and ca. 2 µm shaft diameter). The spermathecal chaetae have a very long, sharply-pointed distal tooth (7–17 µm long, most commonly 12 µm), with a short (3 µm), rounded proximal tooth at the base, nodulus at 0.3 from the distal end (Figs 4 J, 5C). A ventral pad of thickened (to about 30 µm) epidermis surrounds spermathecal chaetae; cells columnar but not distinctly glandular (Fig. 4 K). Modified penial chaetae (72–84 µm long), 2 per bundle (sometimes an additional, partially developed one growing laterally), arranged in parallel, adjacent to male pores in XI, bifid with long, distinct teeth, and nodulus at 0.2 from distal end (Figs 4 I, 5D).</p><p>Nucleated coelomocytes with granulated cytoplasm (diameter 10–22 µm) (Fig. 4 B). Pharyngeal glands in IV (ventro-laterally) and V (ventrally), in some specimens also ventrally in VI. Chloragogenous tissue from the back part of segment IV and densely covering the gut from V backwards. No glands observed behind ventral chaetae. Sperm sac may extend anteriorly to segment VIII and back to XV. Egg sac back to XVI. One pair testes in X, one pair ovaries in XI.</p><p>Spermathecal duct short and bulbous (60–96 µm long, 55–70 µm maximum diameter), formed by a thick wall of columnar epithelium and a muscular layer (Fig. 5 B). Oval spermathecal ampullae occupy most of segment X (188–217 µm maximum diameter). Sperm free in lumen of spermathecal ampulla.</p><p>One pair sperm funnels on 10/11. Vas deferens 20–26 µm wide and about 150–200 µm long, coiled, and joining the atrial ampulla subapically (Fig. 5 B). Atrial ampulla globular (84–111 µm long, 57–81 µm wide), completely covered by a diffuse layer of prostate cells up to 40–57 µm high. Atrial duct (39–45 µm long) shorter than the ampulla and associated with a few muscular strands, its diameter widening from the simple male pore towards the ental end, and not clearly separated from the ampulla (Fig. 5 B). Atrial lumen irregularly filled by epithelial cells (Fig. 4 L).</p><p>Anomalies. In one specimen, genitalia shifted one segment forward.</p><p>Distribution and habitat. Known only from two sites in northwestern Washington. The Calawah River (type locality) and tributaries are low gradient, streams; surface flow varies seasonally, with low flows in July-August. Areas of groundwater upwellings result in localized cold spots. Sediment is dominated by cobble-size stones. Dissolved oxygen levels are high (9.7–10.7 mg /L), and the river is rated Class AA (extraordinary water quality) by the Washington Department of Ecology (Hook 2004). There are no domestic water systems or hatcheries located within the South Fork Calawah, and the watershed has substantial populations of several salmonid species (Hook 2004).</p><p>Remarks. The species is closely related to R. saelonae based on the presence of both penial and spemathecal chaetae, and the prostate gland covering all of the atrial ampulla. It is, however, clearly distinguished from it and other related species (see Table 1) by the smaller number of genital chaetae, the bifid penial chaetae, the structure of the male duct, and the absence of ventral glands in segment X and adjacent segments. The atrial duct is shorter than the ampulla, gradually narrowing towards the male pore in R. quileuticus sp. n., while it is well-separated and longer than the ampulla in R. saelonae .</p><p>Character alcyoneus ardierae carsticus Košel, 1980 gasparoi Martínez- gernikensis Giani &amp; glandulosus (Martínezsp. n. Lafont &amp; Juget, 1993 Ansemil et al. 1997 Rodriguez, 1988 Ansemil et al. 1997)</p><p>Dorsal chaetae Hairs and pectinates. No hairs, bifids Bifids, teeth equal Hairs and pectinates. No hairs, bifids with Hairs and pectinates. Hairs absent from variable (modified in XI) equal teeth Hairs absent from XVII XIII–XV backward backward</p><p>Penial chaetae 5–7 simple-pointed or 3(4–5), simple-pointed, 2–4 bifid, hook-shaped 2 simple or bifid(?) 2 bifid 1–2 simple- pointed</p><p>minutely bifid, fanwise fanwise</p></div>	https://treatment.plazi.org/id/154FD01D4B54FFD8FF6E957EBBD7CE3B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
154FD01D4B5EFFC1FF6E92E6BA3AC9D0.text	154FD01D4B5EFFC1FF6E92E6BA3AC9D0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus clio	<div><p>Rhyacodrilus clio sp. n.</p><p>(Figs 6–8)</p><p>Holotype. USNM 1202056, dissected specimen stained in hematoxylin and mounted in Canada balsam.</p><p>Paratypes. USNM 1202057-61: 3 dissected specimens from type locality (17 July 2011), 2 whole mounts from Squaw Creek at Madrone Camp (18 July 2011). MNCN 16.03/3080: 2 whole mounts from type locality (17 July 2011).</p><p>Type locality. Squaw Creek at Chirpchatter Camp, Shasta Co., California N40.8606° W122.1205° (17 July 2011).</p><p>Other material. 7 mature whole mounts and 1 sagittally sectioned from Santa Clara Co., Coyote Creek, near Gilroy Hot Springs N37.11° W121.47° (22 July 2001, 17 April 2005, 26 March 2010, 16 April 2011). 5 dissected, and 3 whole mounts, stained with hematoxylin, from Shasta Co., Squaw Creek at Madrone Camp, N40.9276° W122.0972° (19 August 1999), 12 whole mounts (18 July 2011). 3 whole mounts, 9 dissected, 3 sagittally sectioned specimens from Squaw Creek at Chirpchatter Camp, N40.8606° W122.1205° (17 July 2011). 5 mature whole-mounts from Napa Co., Napa River drainage, Ritchey Creek, N38.5515° W122.5219° (25 April 2004), collected by R.W. Wisseman. 1 mature whole mount from Santa Cruz Co., San Lorenzo River, approx. N37.02° W122.06° (22 May 1998). 6 mature or nearly mature whole mounts from San Benito Co., Pinnacles National Monument, Chalone Creek near lower park boundary, N36.4598° W121.1549° (27 March 2004). All sites in California (USA).</p><p>Etymology. Named after Clio (Greek Kleio), the Latin name of the muse of history, and thus also of natural history. "To thee, O Muse, has been vouchsafed the power to know the hearts of the gods and the ways by which things come to be" ( Valerius Flaccus, Argonautica 3, 1A.D. (trans. J.H. Mozley). Noun in apposition.</p><p>Description (from specimens of the type locality). Elongated, rounded prostomium (140–200 µm long) (Fig. 6 A). Diameter of the body in segment VIII (whole-mounted specimens) 210–340 µm; maximum diameter at clitellum up to 390 μm. Length of preserved worms 8–15 mm, 56–73 segments. Epidermis non-glandular, 6–10 µm high in anterior segments. Clitellum from level of chaetae in X to XII and weaker in XIII, thickest dorsally and laterally, 16–26 µm high, usually with distinctly glandular epithelium (Fig. 6 B).</p><p>Dorsal bundles composed of 1–3 hairs (commonly 2 in anterior segments, 1 in posterior segments) and 1–3 bifid chaetae. Hair chaetae smooth, usually longer than body diameter in anterior segments, the longest in each bundle 250–450 μm. Dorsal bifid (or pectinate) chaetae (70–86 µm long) in anterior segments (shorter in II), with very fine intermediate teeth (only visible at 1000x), lateral teeth equally long and lyre- or narrowly U-shaped; dorsal bifids shorter and teeth equal in posterior segments, without intermediate teeth (Fig. 6 C, D). Ventral chaetae bifid; anterior ventral chaetae 5–6 per bundle, 70–79 µm long, shorter in II, with distal tooth slightly longer than proximal and about as thick; unmodified ventral chaetae in segment X; posterior ventral chaetae 3–6 per bundle (70–77 µm long) with distal tooth thinner and equal to or shorter than proximal (Fig. 6 E, F). One pair spermathecal pores at the beginning of segment X, in line with ventral chaetae; a pair of transverse (crescent-shaped) shallow grooves (35–45 μm long) just anterior to spermathecal pores, the thicker ends extending mediad, nearly conjoined near the midline, forming a narrow, slightly raised (50–70 μm long, 12–24 μm high) anchorage bridge (Cuadrado &amp; Martínez-Ansemil 2001) between spermathecal pores (Fig. 7 A). One pair male pores in posterior part of segment XI, in line with ventral chaetae. Modified penial chaetae (76–94 µm long) in segment XI, usually 2 per bundle (sometimes with an additional, shorter replacement), although the number of penial chaetae varies among different populations ascribed to the species (see Table 2). Penial chaetae simple-pointed or faintly bifid, and nodulus at 0.2 from distal end.</p><p>Very abundant nucleated and granulated coelomocytes, diameter 12–25 µm (Fig. 6 L). Pharynx short, in II. Pharyngeal glands dorsal and ventral to the gut in IV–V, ventral only in VI; sometimes inconspicuous. Chloragogen layer covering the gut from segment VI. Small clusters of glandular cells on either side of ventral nerve cord; 2–4 pairs per segment beginning at about IV, and continuing through a few post-atrial segments, with narrow cellular extensions through ventral body wall, but without obvious pores (Fig. 6 M). First nephridia visible in VIII in some specimens, or only in postclitellar segments in others; nephridia occur irregularly in postclitellar segments; postseptale granular and usually elongate-narrow; efferent duct and nephropore small and inconspicuous. One pair testes in segment X and one pair ovaries in segment XI. Sperm sac forwards to segment IX and backwards as far as XVIII. Egg sac back to segment XIX.</p><p>Locality Body Hair Dorsal Ventral Penial Spermatheca Maleduct</p><p>diameter chaetae: bifids: chaetae: chaetae:</p><p>in VIII number number number number</p><p>(µm) (length) (length). (length) (length)</p><p>Pectinations</p><p>Squaw 210–340 1–3 (longest 1–3 (70– 86 3–6 (70–79 2 (76–94 µm) Short duct (26–55 µm long), Ampulla (50– 80 x 36–60 µm), a short neck</p><p>Creek to 250–450 µm). Fine µm) shorter One as long as wide (diameter 20–30 µm) and a long, inflated atrial</p><p>µm) intermediate in II specimen duct (70–100 x 55 –75 µm) teeth with 5 on one</p><p>side</p><p>Coyote 203–238 1–2 (150–375 1–3 (46– 65 3–6 (57–71 1 (58–69 µm) Short duct (51–66 µm long), Small globular ampulla (41– 57 x 35–51 µm), a</p><p>Creek µm) µm). No µm) shorter as long as wide short neck (diameter 22–24 µm) and a long, intermediate in II inflated atrial duct (78– 98 x 45–72 µm) teeth</p><p>Spermathecae with a short duct (26–55 µm long, 35–55 µm wide) with high columnar epithelial cells. Spermathecal ampulla large and elongate-sacciform (150–340 µm long, 100–150 μm wide); epithelium in unmated worms irregular, 10–25 μm high (thickest near duct), in mated worms thin (3–6 μm) and more uniform; ampulla usually confined to segment X (Fig. 7 C). Sperm free in the lumen of spermathecal ampulla.</p><p>Paired sperm funnels open in the ventral part of septum 10/11, each leading to a ciliated vas deferens, which joins the atrium at the middle of the ampulla. Vas deferens longer than atrium, length about 200–290 µm, diameter 17–23 µm. Atrium consists of a small globular to ovate ampulla and a long inflated duct; a short neck (20–30 μm long by 30–45 μm wide) connects the ampulla to the atrial duct; this section has narrow columnar epithelium (Fig. 7 B). Atrial ampulla 50–80 μm long by 36–60 μm wide; epithelium granular with large, irregular cells having basal nuclei, lumen narrow (6–7 μm), muscle layer very thin; ampulla covered by a diffuse layer of prostate cells, 20–35 µm high, usually a mix of small clusters and individual cells. The outer duct forms an eversible penis; entire duct 70–100 μm long by 55–75 μm wide when penis is retracted, with thick, granular epithelium (10–14 μm) having basal nuclei and a thin (1–2 μm) outer muscle layer. Everted penis short and broad, formed by lining cells from outer part of atrial duct extending through pore (Figs 6 K, 7D, E). Penial chaetae terminate within a small tubercle at the inner margin of the male pore, with a small patch of glandular cells, and several muscle strands extending to dorsal body wall (Fig. 7 B, C).</p><p>Distribution and habitat. Known from several localities in central and northern California. Sites were small to mid-sized, gravel-cobble bed streams in relatively undisturbed, foothill settings; worms were collected from riffle habitats with coarse sediments. Sites on Coyote and Chalone creeks are intermittent, typically losing surface flow in summer.</p><p>Remarks. Rhyacodrilus clio sp. n. has been ascribed to the genus Rhyacodrilus based on the diffuse prostate layer covering the atrial ampulla, the vas deferens, which is longer than the atrium and joins it subapically, and the presence of modified penial chaetae. The new species is well-separated from other Rhyacodrilus species by the structure of the atrium, with a small ampulla and a large inflated duct that can be partially everted, with the inner epithelial cells of the duct extending beyond the male pore. Different populations that have been ascribed to R. clio show different numbers of penial chaetae and some variations in the range of measurements of chaetae and atrium (see Table 2). However, the general structure of male duct and spermatheca are consistent (Fig. 8). The individuals from Ritchey Creek were collected in alcohol, and thus poorly preserved; they have been provisionally ascribed to this species based on shape of chaetae, the short atrial ampulla, and the short, everted penes.</p><p>Rhyacodrilus clio sp. n. appears to be related to the European groundwater species Rhyacodrilus subterraneus Hrabë, 1963 b, with which it shares the presence of hair and pectinate chaetae in dorsal bundles and penial chaetae but no spermathecal chaetae. As in R. clio sp. n., the atrium of R. subterraneus is composed of a small ampulla covered by diffuse prostate glands and a long duct; although the duct is thinner, it has been described as having cubic epithelium, and appears to be devoid of a penis. The shape of both dorsal and ventral anterior chaetae also distinguishes both species: R. clio has lyre or U-shaped pectinates, with teeth of equal length, while R. subterraneus has unequal V-shaped pectinates, and anterior ventral chaetae with upper tooth much longer than lower.</p><p>Rhyacodrilus subterraneus has been reported in several parts of Europe (Giani &amp; Martínez-Ansemil 1981, Timm et al. 1996, Martin et al. 2009, Schenková et al. 2010), although some of these records may need to be reconsidered. A re-examination of the specimen reported by Giani and Martínez-Ansemil (1981) from a NW Spanish locality revealed the typical chaetal morphology of R. subterraneus (Fig. 9 K, L). However, this specimen is probably a different species, as the atrium has a large ampulla relative to the short, narrow atrial duct. Timm et al. (1996) reported R. subterraneus from several locations in the Scandinavian Peninsula, but the studied specimens may be uncompletely mature since the spermathecal ampulla is drawn empty, and the atrium does not show an internal lumen nor a penis. Rhyacodrilus subterraneus has also been reported from several localities in eastern and central United States (Strayer &amp; O'Donnell 1988, Strayer 2001, Wetzel &amp; Taylor 2001, Wetzel &amp; Morgan 2007), but these records must also be interpreted with caution, as they are based on chaetal characters of immature specimens (see also remarks for R. propiporus sp. n. below).</p></div>	https://treatment.plazi.org/id/154FD01D4B5EFFC1FF6E92E6BA3AC9D0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
154FD01D4B41FFC4FF6E9766BD04CCDB.text	154FD01D4B41FFC4FF6E9766BD04CCDB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus propiporus	<div><p>Rhyacodrilus propiporus sp. n.</p><p>(Figs 9–11)</p><p>Holotype. USNM 1202062, whole-mounted worm stained in hematoxylin and mounted in Canada balsam.</p><p>Paratypes. USNM 1202063-65: 1 dissected specimen (14 February 2007), 1 sagittally sectioned (17 March 2007), 1 whole mount (17 March 2007), MNCN 16.03/3081: 1 dissected specimen (17 March 2007), all from the type locality.</p><p>Type locality. Beech Swamp, Halifax Co., North Carolina, USA, N36.1435° W77.5551° (17 March 2007).</p><p>Other material. 2 dissected and 1 whole mount from Lower Little River at road SR 2031, Harnett Co., N35.2599° W78.8230°, (5 June 2011); 4 dissected, 2 whole mounts, 1 sagittally sectioned (July 2011). 2 whole mounts from Lower Little River at highway NC 217, N35.2637° W78.7406°, (5 August 2011). 6 whole mounts from Lower Little River at highway NC 87, Cumberland Co., N35.4001° W79.1126° (5 August 2011). 2 dissected from Powell's Creek at road SR 1338, Halifax Co., N36.1791° W77.8980° (13 December 2011). 1 whole mount from Friar Swamp at road SR 1740, Columbus Co., N34.3695° W78.4572° (1 March 1998). 1 whole mount from seep at Pettiford Creek, Carteret Co., approx. N34.75° W77.02° (30 January 2009). 1 whole mount from Chinkapin Creek, Hertford Co., N36.2534° W76.8491° (1 March 2001). All material collected by David R. Lenat from North Carolina, USA.</p><p>Etymology. From the Latin prope for close and porus for pores. The specific name refers to the very close (nearly median) position of both spermathecal and male pores.</p><p>Description (from type locality specimens). Number of segments 63–74. Body diameter 440–456 µm in segment VIII, 556–614 µm in XI. Rounded-conical prostomium 107–202 µm long. Epidermis 7–13 µm high, 25– 45 µm at clitellum. Clitellum from the level of chaetae in X to the end of XII.</p><p>Dorsal bundles in anterior segments with 0–3 smooth hair chaetae (229–242 µm long) and 3–4 bifid, nonpectinate chaetae (103–107 µm long) with distal tooth longer than proximal (Fig. 9 B–E). Hair chaetae absent from segment VIII (or IX) backwards. Posterior dorsal chaetae like ventral chaetae (69–110 µm long). Ventral chaetae bifid; anterior ventral chaetae 4–6 per bundle (73–110 µm long) with distal tooth longer than proximal; posterior ventral chaetae 2–4 (commonly 3) per bundle, 88–99 µm long, and progressively shorter, down to 70–74 µm long in the most posterior segments of complete individuals, distal tooth becoming equal to or shorter than proximal in the most posterior segments (Fig. 9 F–I).</p><p>One pair spermathecal pores at the beginning of segment X, close to the mid-ventral line of the body, usually within a shallow, common groove about 150 μm wide; the spermathecal pores surrounded by a thin, muscular ring at the body wall (similar to Fig. 11 C). One pair male pores at about the posterior 1/4 of segment XI, close together near the mid-ventral line, on a prominent porophore 80–120 μm wide and up to 45 μm high, lined with a thin, circular muscle layer (Figs 9 M, N, 10A). No modified genital chaetae. Ventral chaetae usually absent in X at maturity; in one mated worm a single chaeta was not modified and resembled other ventral chaetae (79 µm long). Female pores in intersegment 11/12, just below lateral line.</p><p>Abundant nucleated coelomocytes (diameter 10–24 µm), containing large granules (Fig. 9 A). Pharyngeal glands in V and poorly developed in IV and VI. Chloragogenous tissue scantily covering the gut from VII backwards, and formed by a thin layer of cells (usually &lt;20 µm high). One pair testes in segment X and one pair ovaries in segment XI. Sperm sac back to segment XV, egg sac back to segment XVI. Spermathecae composed of a short duct (43–77 µm long and 58–64 µm maximum diameter) and an ampulla (154–246 µm long, 138–215 µm wide) in the most anterior part of segment X (Fig. 10 B, C). Duct lined with columnar epithelium, with a narrow (5– 10 μm) lumen and very thin muscle layer. Ampulla with thin layer of cuboidal epithelium (6–21 μm); lumen with sperm typically arranged in loose clusters.</p><p>Male duct consisting of a sperm funnel at the most ventral part of septum 10/11, and a relatively straight vas deferens which joins the atrial ampulla subapically; vas deferens of homogeneous width (diameter 23–24 µm), longer than atrium (ca. 230–315 µm long) (Fig. 10 B). Atria formed by a spherical ampulla (diameter 57–75 µm) and a duct (78–125 µm long, 43 µm maximum diameter) about 1.5–2 times longer than the ampulla; duct tapering towards the male pore and usually towards the ampulla too. Penis absent. Atrial ampulla with a thin epithelium (7– 8 µm) and very thin musculature, covered by a diffuse layer of prostate cells (up to 10–14 µm high), not forming bundles of cells (Fig. 10 D–F). The relative size of the atrium is very small, total length less than one third the diameter of the body.</p><p>Lower Little River population (supplemental description, differences from type locality population). Body dimensions generally smaller than Beech Swamp population (type locality): number of segments 53–72, diameter 270–400 µm in segment VIII and 360–490 µm in XI (slide-mounted worms). Chaetae similar, but usually shorter than in Beech Swamp population: anterior dorsal bundles with 1–3 smooth hair chaetae (maximum 200–260 µm long, shortest in II and X) and 3–4 bifid chaetae (85–110 µm long) with distal tooth slightly or up to 2 times longer than proximal. Hair chaetae absent from segment X–XII backwards. Posterior dorsal chaetae like ventral chaetae (both 85–100 µm long). Anterior ventral chaetae 4–8 per bundle (95–115 µm long) with distal tooth longer than proximal (sometimes with a thin intermediate tooth). Posterior to clitellum 2–5 ventral chaetae; commonly 3 per bundle in posterior half of worm, with distal tooth becoming equal or shorter than proximal in the most posterior segments.</p><p>Spermathecal pores usually within a shallow, common groove 50–70 μm wide, pores surrounded by a thin, circular muscle layer ectal to the ventral muscle bands of the body wall (Fig. 11 C). Male pores on a low porophore about 180–240 μm wide, and up to about 40 μm high; porophore lined with a thin, circular muscle layer.</p><p>Spermathecal duct may be cylindrical and up to twice as long as wide, or short and conical, about as wide as long entally (Fig. 11 A, B) (length 55–96 µm, width 38–62 µm). Spermathecal ampulla ovate (125–245 µm long, 100–180 µm wide in mated worms), usually entirely within X. Atrial ampulla spherical to slightly ovate (60–145 µm wide); duct cylindrical or tapered (55–104 µm long, 30–42 µm maximum diameter); total length of atrium 120–225 µm, about one third to half the diameter of the body at segment XI; relative lengths of duct and ampulla variable, but usually about equal (Fig. 11 A, B). Atrial ampulla with epithelium 10–20 µm high, very thin musculature, and covered by a diffuse (usually single) layer of prostate cells.</p><p>The other listed populations from different locations in North Carolina are represented by limited material, and have been provisionally ascribed to R. propiporus . Specimens from Powell's Creek had very similar reproductive organs, but were unmated and had unmodified ventral chaetae in segment XI (Fig. 11 D).</p><p>Anomalies. One Lower Little River specimen had paired spermathecae in VII and VIII, paired atria in VIII, and a single atrium in IX; only the second male duct had sperm on the funnel. Another specimen had normal reproductive organs, but they were shifted forward by two segments. Six mated specimens had 1 or 2 unmodified chaetae in XI, but these were presumably remnant somatic chaetae, as they were on the line of ventral chaetae, and were not medially directed.</p><p>Distribution and habitat. Rhyacodrilus propiporus sp. n. was collected in diverse habitats in eastern North Carolina. Beech Swamp (the type locality) and Powell's Creek are in the Tar River basin, in the northeastern part of the State. Rhyacodrilus habitats were small (2 m wide), sandy streams that typically go dry during summer months. Both streams appear to have good water quality based on the macroinvertebrate composition. Beech Swamp has a diverse oligochaete community, and is the only known location for the monotypic Pilaridrilus Fend and Lenat, 2007 (Lumbriculidae) .</p><p>The Little River is in southeastern North Carolina, in the Cape Fear River basin. It is a large (20 m wide), permanent stream, with low gradient and excellent water quality based on the macroinvertebrate community (D.R. Lenat, pers. comm. 2012). Rhyacodrilus propiporus was usually collected from sand and gravel near the middle of the river.</p><p>Remarks. Rhyacodrilus propiporus sp. n. belongs to a small group of Rhyacodrilus species with hair chaetae in dorsal bundles and a layer of prostate cells covering the atrial ampulla, and lacking both modified spermathecal and penial chaetae. This group is formed by R. lepnevae Malevich, 1949 emend. Hrabë (1974) (Russia), and R. lindbergi Hrabë, 1963 a (described from one immature specimen from a cave in Portugal). Martínez-Ansemil et al. (1997) attributed a single specimen with penial chaetae from northwestern Spain to R. lindbergi, although this character was not verified in the type specimen (E. Martínez-Ansemil, pers. comm. 2011). Both species are distinguished externally from R. propiporus sp. n. by the presence of ventral bundles with both simple-pointed and bifid chaetae. Externally, the near-median positions of the spermathecal and male pores distinguish the new species from other Nearctic members of the genus. The new species is also well-separated from other Nearctic Rhyacodrilus by the small size of the atrium and relative lengths of the atrial ampulla and duct (the latter being 1.5 to 2 times the length of the former), and also by the very short, one cell layer prostate covering the ampulla (unique among the Nearctic Rhyacodrilus species).</p><p>The population from Lower Little River has been provisionally ascribed to R. propiporus based on similar chaetal characteristics and the peculiar near-median position of male and spermathecal pores. The larger atrium, a relative length of atrial ampulla to duct of about 1, a cylindrical spermathecal duct, and ventral chaetae often present in segment X, distinguish this population from that of the type locality, and its status may be subject to revision with new findings in eastern North America.</p><p>This species may have a broader distribution than indicated here. The combination of dorsal hairs limited to anterior segments and anterior dorsal bifid chaetae with the distal tooth longer than the proximal would likely attribute immature R. propiporus to R. subterraneus using the keys by Kathman and Brinkhurst (1998).</p><p>Examination of material collected by D. Strayer from several sites in eastern USA, and cited by Kathman and Brinkhurst (1998) as R. subterraneus, revealed no mature specimens, so the presence/absence of penial chaetae cannot be verified. All 6 individuals examined from the USNM collection (# 102832, #187041 and #187042, leg. Strayer), plus 2 additional specimens from the Strayer collection, were immature, with upper tooth of dorsal chaetae thicker and much longer than lower (as in R. subterraneus), and with short intermediate teeth visible in some individuals but not in others. In the absence of mature worms attributable to R. subterraneus, we conclude that at least some of the North American records of R. subterraneus based on chaetal morphology may be R. propiporus, and that an examination of the male ducts and/or molecular analyses are required to confirm the presence of R. subterraneus in North America (see also remarks for R. clio sp. n. above).</p></div>	https://treatment.plazi.org/id/154FD01D4B41FFC4FF6E9766BD04CCDB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
154FD01D4B44FFC8FF6E9068B8EFCE00.text	154FD01D4B44FFC8FF6E9068B8EFCE00.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus alcyoneus	<div><p>Rhyacodrilus alcyoneus sp. n.</p><p>(Figs 12–13)</p><p>Holotype. USNM 1202066, mature but unmated specimen, dissected and stained in hematoxylin; mounted in Canada balsam.</p><p>Paratypes. USNM 1202067-69, 1 dissected specimen stained in hematoxylin, from the type locality (20 June 2011). 2 partially mature, dissected worms from the type locality (7 June 2008 and 25 March 2007).</p><p>Type locality. Guadalupe Creek, above Guadalupe Reservoir, in a short (ca. 50 m) reach below the confluence with Rincon Creek, N37.1831° W121.8716°, Santa Clara Co. California (USA) (21 June 2011).</p><p>Other material. From the type locality. Slide-mounted, partially mature specimens: 6 dissected (25 March 2007, 6 April 2008, 11 May 2009, 7 June 2008). 20 unmounted, partially mature specimens: 2 from 22 March 2008, 4 from 6 April 2008, 3 from 17 April 2008, 10 from 27 March 2009, 1 from 8 March 2010.</p><p>Etymology. alcyoneus, for Alcyoneus, the eldest of the gigantes ("the earth born" in Greek) slain by Heracles. The name refers to the very large size of the worms in this species, relative to congeners.</p><p>Description. Whole body dimensions in the description are based on 24 worms, all with at least obvious sperm sacs or partially developed genital pores. Number of segments in complete specimens about 100–135, length up to 63 mm (most specimens about half this length; median 41 mm). Diameter of the body in VIII 0.9–1.3 mm; maximum diameter to 1.6 mm. Short, round prostomium (315–486 µm long, 504–630 µm wide at base) (Fig. 12 A). Epidermis glandular, 15–33 µm high in anteclitellar region, where it is more darkly stained in haematoxylin than in postclitellar region. Secondary annulation usually visible in anterior segments as a narrow anterior ring; median and posterior furrows often present, dividing each anterior segment into 2–4 apparent sections (Fig. 12 A, B); apparent segmentation in posterior segments varies with fixation, but additional rings may be visible. Mid-dorsal pores at the anterior secondary furrow in III–VII (Fig. 12 B, E).</p><p>Clitellum saddle-shaped (thickest dorsally), maximum thickness 70–100 μm, from level of chaetae in X to mid-XIII. One pair spermathecal pores at the beginning of segment X, slightly lateral to the line of ventral chaetae, on the secondary annulation; actual pore is at the outer edge of an inconspicuous transverse groove (Fig. 13 B). One pair male pores in segment XI, in line of ventral chaetae, externally visible as transverse folds or pits. One pair female pores, inconspicuous at 11/12, about midway between the lateral line and the line of ventral chaetae.</p><p>Dorsal bundles in anterior segments with 1–3 smooth hair chaetae (maximum length 530–764 µm, shorter in segment II up to 240–370 µm) and (1)3–6 pectinate chaetae. Pectinate chaetae up to 124 µm long in II, 166–224 µm in other segments, very slightly sigmoid, teeth about equally long, and pectinations shorter than or equal to outer teeth (Fig. 12 F, G). Posterior dorsal chaetae 3–6 per bundle, not pectinate, much like ventral chaetae posterior to about XV (155–237 µm long), upper tooth thinner and of the same length or slightly shorter than lower (Fig. 12 H). Hair chaetae absent posterior to segment XIII or XIV. Ventral chaetae bifid. Anterior ventral chaetae 4–6 per bundle (141–200 µm long), sigmoid, with nodulus about 1/3 the distance from the tip; distal tooth 1.5 to 2 times the length of proximal. In mature worms ventral chaetal bundles of IX and X on low, round tubercles, 170–220 μm wide (Fig. 13 B cht); ventral chaetae in X reduced to 2(3) per bundle, slightly longer (245–288 μm) than those in anterior segments, and with distal tooth up to 4 times longer than proximal (Fig. 12 M). Postclitellar ventral chaetae 1–4(6), commonly 2–3, per bundle (133–230 µm long); teeth about equal in length, distal tooth thinner. Modified penial chaetae unequal in length (221–327 µm) in segment XI, 5–7 per bundle, simple-pointed or minutely bifid in the most mature specimens (or bifid in some of the less-mature worms), and arranged fanwise at male pore (Fig. 12 K, L).</p><p>Very abundant nucleated and granulated coelomocytes (diameter 24–48 µm) (Fig. 12 C). Pharynx in II–III (IV) with a well-developed dorsal pad of glandular tissue interspersed with muscle strands. Pharyngeal glands from IV to V, or restricted to IV. Chloragogen layer covering the gut from segment V backwards, up to 108 µm high, densely granulated (Fig. 12 D).</p><p>Descriptions of reproductive organs are based on dissections of 3 unmated specimens with mature eggs in the egg sacs, collected on 20–21 June 2011, and supplemented with less-mature material collected on other dates. One pair testes in segment X and one pair ovaries in segment XI. Large eggs with yolk granules visible in 3 individuals, egg sac to XX–XXVI; sperm sac back to segment XVIII–XXIV.</p><p>Spermathecal duct joins a wide ectal vestibule at lateral edge of outer fold (135–175 μm long); the duct is very short (45 μm long by 100–175 μm wide), with narrow-columnar epithelium and a very narrow lumen (Fig. 13 A, B). Spermathecal ampulla without sperm in all examined individuals (i.e. unmated), up to 340 by 490 μm.</p><p>Sperm funnel opens in the ventral part of septum 10/11, up to 300 μm wide. Vas deferens ciliated, with columnar epithelium in middle portion; diameter 51–66 µm, narrowing to 27–45 µm close to the atrial junction, and to 30–54 µm at the sperm funnel. Vas deferens longer (520–700 µm) than atrium, joining atrium subapically (Fig. 13 A, B). Atrium elongated and roughly tubular, narrowed at ends, 405–510 μm long in the 3 most-mature worms, 360–469 µm in the other material, and 82–125 µm wide. No clear separation between atrial ampulla and duct. Ental 1/2 to 2/3 of atrium ampulla-like, with somewhat columnar epithelium, lumen ciliated, muscle layer about 3 μm thick and covered by a layer of prostate cells (70–120 µm high) in tightly-packed bundles in more mature worms, or appearing diffuse in less-mature worms. Ectal 1/2 to 1/3 of atrium without prostate, 180–220 μm long, to 122 μm wide entally, narrowing to 78 μm ectally; with oblique muscle strands extending from the atrial muscle layer (5 μm) to the body wall near the male pore. A mass of loosely packed cells (possibly glandular) among the muscle bands, associated with the male pore. Several dorso-ventral muscle strands are associated with the bundles of penial chaetae at the male pores.</p><p>Distribution and habitat. Guadalupe Creek occupies a small drainage in the Coast Ranges of central California. The stream is regulated, but the alcyoneus type locality is upstream of the reservoirs, and has a relatively undisturbed watershed with natural flow regime. The collection site is a third-order stream with gravelcobble sediments; summer temperatures are generally low (below 20° C), and salmonids are present (Carter &amp; Fend 2000; Jae Abel, Santa Clara Valley Water District, pers. comm. 2012). Surface flow is very low from late summer through autumn, and is intermittent in some years. The section of the river Guadalupe where the population has been sampled has localized sulfide upwellings.</p><p>Remarks. The size of Rhyacodrilus alcyoneus sp. n. relative to other Nearctic Rhyacodrilus is one of the most striking characteristics of this species. Two other very large Rhyacodrilus species, R. intermedius Semernoy, 2004 and R. multispinus (Michaelsen, 1905), have been reported from Lake Baikal, although both of them are devoid of hair chaetae. The former has been reported as 50 mm long by 1.5 mm wide (Semernoy 2004) and the latter up to 57 mm long by 4.0 mm wide (Chekanovskaya 1962). The extremely large dimensions (80-150 mm long) reported in the description of R. sinicus Chen, 1940 (and repeated in the literature, e.g. Ohtaka 1995) are surely an error in the original description, as Chen (1940) also stated that the species is similar in size to other tubificids, the number of segments (42–56) is quite modest, and and hair chaetae are stated to be about the diameter of the body (240–250 μm).</p><p>All examined individuals are from the type locality in Guadalupe Creek. Collections were made on several dates between 2007 and 2011. Although sampling was done at other times during the year, Rhyacodrilus alcyoneus sp. n. has only been collected from the period of January to June. The reproductive organs appear completely developed in only 3 specimens of the available material. These individuals had mature eggs with yolk granules in the egg sac, but all of them lacked sperm in the spermathecal ampulla and in the sperm funnel, suggesting that this population fails to reproduce sexually and is parthenogenetic. According to Edwards and Bohlen (1996), the association between parthenogenesis and high polyploidy in earthworms is an advantageous condition that provides resistance to environmental stress (Cosin et al. 2011).</p><p>The structure of the male duct is very similar to that described below for R. longichaeta sp. n. found at another site on the same stream. However, R. alcyoneus sp. n. never develops the characteristic long hairs in segment II that are diagnostic for R. longichaeta . Other characters that consistently distinguish R. alcyoneus sp. n. from R. longichaeta are the lack of hair chaetae in postclitellar segments, the posterior dorsal chaetae resembling the ventral chaetae, the distinct spermathecal vestibule, the larger number (more than 5) of penial chaetae, the large chaetal tubercles in IX and X, and the pharyngeal glands not extending posterior to segment IV or V.</p><p>Two Siberian species, Rhyacodrilus brevis Semernoy, 2004 and R. sibiricus Semernoy, 1971, also have hair chaetae restricted to anterior segments, although atria of these species are more or less globular instead of tubular. Based on the presence of hair chaetae in dorsal bundles, modified penial and spermathecal chaetae and a tubular atrium, R. alcyoneus sp. n. is closer to R. svetlovi Sokolskaya, 1976 from the Chukchi Peninsula (East Siberia), but the vas deferens in that species is shorter than the atrium, and joins it apically instead of subapically. Compared with other Rhyacodrilus species having modified penial chaetae and tubular atria but unmodified spermathecal chaetae, R. alcyoneus sp. n. is well-distinguished by several features. The subapical junction of the vas deferens with the atrium distinguishes R. alcyoneus from R. korjakovi Semernoy, 2004, R. vasalatus Semernoy, 2004 and R. suputensis Timm, 1990, where the vas deferens joins the atrium apically. It is also well-separated from the Baikalian species R. korjakovi and R. vasalatus by the absence of a clearly separated atrial duct. It is also worth mentioning that R. korjakovi loses the hair chaetae when it matures (while keeping the pectinates), and R. vasalatus only has bifids.</p><p>Among Rhyacodrilus species, the modified spemathecal chaetae are usually very different from somatic ventral chaetae (i.e. long or grooved distal teeth, like in R. saelonae sp. n. or R. carsticus Košel, 1980); however, in R. alcyoneus sp. n. they are larger with longer distal teeth, but not very different in shape.</p><p>The presence of dorsal pores in this species is an interesting feature, since after Caramelo and Martínez- Ansemil (2012) the only microdriles (apart from enchytraeids) known to have dorsal pores belong to the subfamily Rhyacodrilinae: Monopylephorus aucklandicus (Benham, 1909), Bothrioneurum vejdovskyanum Štolc, 1886, Peristodrilus montanus, and Protuberodrilus tourenqui . Stephenson (1930) described their function in earthworms as related to the protection against desiccation, keeping the body wall moist for gas exchange, and providing protection against bacteria and parasites. The first pore is in the secondary annulation, at the front of segment III, while those reported in other rhyacodrilines were located in the intersegments 3/4 or 5/6 (Caramelo &amp; Martínez- Ansemil 2012: Fig. 5).</p></div>	https://treatment.plazi.org/id/154FD01D4B44FFC8FF6E9068B8EFCE00	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
154FD01D4B48FFF7FF6E9116BB80CF03.text	154FD01D4B48FFF7FF6E9116BB80CF03.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Rhyacodrilus longichaeta	<div><p>Rhyacodrilus longichaeta sp. n.</p><p>(Figs 14–16)</p><p>Holotype. USNM 1202070, mature worm with sperm in the spermathecae, stained in hematoxylin, dissected and slide-mounted in Canada balsam.</p><p>Paratypes. USNM 1202071-74, 4 mature worms (2 with sperm in the spermathecae): 2 sagittally sectioned, 2 dissected and slide-mounted (23 June 2011); MNCN 16.03/3082: 1 whole mounted, unmated worm (15 May 1998), 1 dissected worm (6 April 2008). All from type locality.</p><p>Type locality. Guadalupe Creek, above Guadalupe reservoir, N37.1822° W121.8725°, Santa Clara Co., California (23 June 2011).</p><p>Other material. 18 partially mature specimens from the type locality: 2 mature unmated individuals, with sperm on male funnels, one sectioned (23 June 2011) and one dissected (20 June 2011). Other individuals with well-developed reproductive organs, with visible sperm sacs but sperm not fully developed: 2 whole mounts (15 May 1998); 5 dissected, (2 from 19 March 1999, and 2 from 9 April 2011); 8 whole mounts (25 April 2006). Additional unmounted worms from the type locality, all partially mature with visible sperm sacs: 6 in alcohol (19 March 1999), 14 in alcohol (25 April 2006). Immature specimens tentatively attributed to R. longichaeta: 7 immature whole mounts from Cooper River, above Cooper Lake, N47.4357° W121.1911°, Kittitas Co., Washington (1 November 1990); 3 immature whole mounts from the Merced River in Yosemite Valley, N37.7407° W119.5666°, Mariposa Co., California (6 October 1994 and 16 September 1995).</p><p>Futher material studied (not included in the description). Type series of Edmondsonia montana Brinkhurst, 1965: Holotype USNM 32652 (1 slide), Paratype USNM 32653 (9 slides), all from Dog Lake, Yakima Co. Washington. An additional specimen attributed to this species, USNM 32980, from Sugar Island, Michigan.</p><p>Etymology. The specific name, longichaeta, refers to the extraordinarily long hair chaetae in segment II.</p><p>Description (from type locality specimens). Number of segments 48–95. Prostomium rounded to conical, 180–360 µm long (Fig. 14 A). Body diameter 0.42–0.74 mm in VIII. Secondary annulation usually absent in anterior segments, or a weak anterior ring in about IX–XI; variable in posterior segments, with 0–3 narrow rings. Mid-dorsal pores in intersegments (3/4) 4/5 to 6/7 (7/8). One pair spermathecal pores in the most anterior part of segment X, in line with ventral chaetae; two crescent-shaped grooves demarcate a median raised area between the pores, interpreted as a copulatory anchorage bridge. One pair male pores in the line of ventral chaetae in segment XI, usually opening on small round papillae; modified penial chaetae located closer to the body midline (Fig. 14 C). One pair female pores at intersegment 11/12, between lateral line and line of ventral chaetae.</p><p>Faintly hispid hair chaetae 1–4 per bundle in anterior segments (170–540 µm long), in segment II 1.4 to 2 times (620–960 µm) longer than the longest hairs in other preclitellar segments (Figs 14 A, 15F); hair chaetae present in most postclitellar segments, 0–1 per bundle, length less than 200 µm. Lateral hairs of hispid chaetae difficult to see in specimens from Guadalupe Creek (1000x usually required), but clearly visible at 400x magnification in Cooper and Merced River populations (Fig. 15 P). In anterior segments, 3–5 pectinate chaetae per bundle (96–144 µm long), with 4–7 intermediate teeth shorter than lateral teeth; 2–3 per bundle in posterior segments, commonly with very fine pectinations visible only at 1000x (Fig. 15 E–H). Upper teeth of anterior pectinates are slightly longer and thinner than, or equal to lower, although this varies even within the same bundle (Fig. 15 E); differences in size between lateral teeth are not as marked as in other populations (e.g. Cooper and Merced Rivers), or as in the type series of R. montana (Fig. 15 O, 16A, D, E). Ventral chaetae bifid; in anterior bundles 4–6(8) (96–149 µm long), with distal tooth thick and 1.5–2.5 times longer than proximal (longest in the most lateral chaetae within each bundle); 3–5 ventral chaetae in posterior segments (110–115 µm long) with distal tooth as long as but thinner than proximal (Fig. 15 I–M). In mature worms with sperm on the male funnels, ventral chaetae in segment X reduced in number (2–3 per bundle) and slightly larger (150–185 µm long), 1.2–1.8 times longer than chaetae in IX; the shape is also slightly modified, with distal tooth up to 3 times longer than lower (Fig. 14 B, 15N). At the male pore, (2)3(4) penial chaetae arranged more or less fanwise, simple-pointed or bifid (Fig. 15 D), 150–176 µm long, and associated with 2 strong dorso-ventral muscular strands (Fig. 14 E).</p><p>Pharynx with a well-developed dorsal muscular pad. Chloragogen cells start in segment VI and form a layer up to 27 µm high. Large coelomocytes (diameter 21–39 µm), nucleated and granulated (Fig. 15 A). Epidermis 11–18 µm high, circular muscle layer 5–8 µm thick and longitudinal muscles 10–30 µm thick. Clitellum 45–54 µm high dorsally, usually from about 1/ 2 X –XIII, may extend anteriad as far as the line of chaetae in IX. Pharyngeal glands from IV to VI, VII or VIII, in one individual back to the anterior part of IX.</p><p>Sperm funnel on septum 10/11. Vas deferens ciliated (ca. 390–750 µm long, 30–50 µm wide), narrowing (18– 30 µm) before subapical junction with atrium. Atrium club-shaped, total length 250–350 µm. Atrial ampulla elongate (length 110–168 µm, maximum diameter 54–74 µm), covered by a diffuse layer of prostate cells (up to 28 µm high), with thin musculature (ca. 3 µm thick). Atrial duct (96–192 µm long, 30–53 µm wide) not separated from ampulla by any constriction, although clearly distinguished by the absence of prostate cells, high columnar epithelium and prominent muscle strands that externally attach it to the ventral region around the male pores (Fig. 14 C, D, F). Sperm sac back to segments XVI–XXVII, egg sac back to XVI–XXVII.</p><p>Spermatheca with a short duct (55–108 µm long, 62–73 µm wide) that opens at the beginning of segment X, in the line of ventral chaetae. Spermathecal ampulla 168–305 µm long, of irregular sac-like shape, with unordered sperm (Fig. 14 C).</p><p>Anomalies. One individual from Guadalupe Creek had the genitalia shifted 3 segments forward (male pores in VIII) suggesting regeneration after fragmentation; one other specimen appeared to have a regenerating prostomium.</p><p>Distribution and habitat. Within Guadalupe Creek, R. longichaeta was found only in gravel-cobble sediments in riffle habitats above the confluence with Rincon Creek. Rhyacodrilus alcyoneus sp. n. was found only in riffles below this confluence, and neither has been found in downstream Guadalupe Creek sites, below the major reservoir. A benthic macroinvertebrate survey of Santa Clara Valley streams (Carter &amp; Fend 2000) did not produce any other records of these species.</p><p>Remarks. Rhyacodrilus longichaeta sp. n. belongs to the group of Rhyacodrilus species with both hairs and pectinates, and also with several penial chaetae arranged fanwise and modified spermathecal chaetae. Within this group, R. svetlovi Sokolskaya, 1976 and R. alcyoneus sp. n. have a similar elongate, tubular atrium, with the atrial ampulla not separated from the duct by a constriction. The vas deferens of R. svetlovi is shorter than the atrium, and joins it apically. Differences from R. alcyoneus (described in the present paper) include the smaller size, the chaetal characteristics, the absence of ventral chaetal tubercula in IX and X, and the presence of hair chaetae in dorsal bundles of postclitellar segments. Both species have similar male ducts, with elongate-tubular atria and a weakly differentiated atrial duct; differences are not very pronounced, with the atrial duct less histologically differentiated in R. alcyoneus, and the simple male pore on a round papilla in R. longichaeta but opening within a fold in R. alcyoneus . Other Rhyacodrilus species with modified penial chaetae (but without spermathecal chaetae) and clubshaped to tubular atria are R. vasalatus and R. korjakovi, described by Semernoy (2004) from Lake Baikal. Of these two species, the male duct in R. longichaeta resembles that of R. vasalatus in having a vas deferens which narrows considerably before the atrial junction, and also in the high columnar epithelium of the atrial duct. However, R. vasalatus has only bifid chaetae, as does R. korjakovi when it is mature.</p><p>The lateral position of the female pores is uncommon in oligochaetes, where they typically have a more ventral position, in line with the ventral chaetae. Interestingly, this character is shared with other western American Rhyacodrilus species described here, i.e. R. saelonae, R. clio and R. alcyoneus .</p><p>Rhyacodrilus species with long hairs in II have generally been attributed to Rhyacodrilus (= Edmondsonia) montana (Brinkhurst, 1965), although elongate hairs in II were not mentioned in the original description. Furthermore, although diagnostic characters in Rhyacodrilus are largely based on details of the reproductive system, the original description of R. montana was based on immature worms, and only included characteristics such as the number and shape of chaetae and the abundance of coelomocytes (see Fig. 16). The type series was collected at three sites in western North America: two from the northern Cascades Range in Washington (Dog Lake and Lake Tipsoo) and a distant California site (a pool at the edge of the Kings River); the holotype and a large series of paratypes were from Dog Lake. The species was transferred from the invalidated genus Edmondsonia Brinkhurst, 1965 to Rhyacodrilus by Brinkhurst and Cook (1966), based on the study of several populations of the Great Lakes region (eastern North America), but in that publication there is no comparison of chaetal characteristics with the type series and only limited information on the male duct.</p><p>Our examination of the R. montana holotype (USNM 32652) and all paratypes (USNM 32653) from the type locality has confirmed that the worms are immature. Hairs are not elongate in segment II of the holotype (485 µm in II and 345–530 µm in the following preclitellar segments) and of most paratypes. However, they are slightly longer in 3 paratypes (600–725 µm in II, and 500–650 µm in the following preclitellar segments, ratio of 1.1 between the longest in II and the longest in following segments); additionally, hairs are smooth, although they are originally described as "feathered hairs" (Brinkhurst 1965: 155). The anterior pectinates in R. montana have the distal tooth longer than the proximal (2 times longer in segments II–VIII of the holotype, about equal posteriorly), and ventral chaetae are pectinate (Fig. 16 B). The poor state of conservation of the tissues does not permit observations on internal anatomy, and even the chaetae are sometimes difficult to see. Accordingly, we propose that R. montana be considered species inquirenda due to its doubtful identity.</p><p>The long hairs in segment II are an obvious feature for species identification, since they can be clearly seen under the stereoscopic microscope. These chaetae, and the fanwise arrangement of the penial chaetae, were first reported for worms attributed to R. montana from the Great Lakes region by Brinkhurst and Cook (1966: 23) and used as diagnostic characters in subsequent literature (e.g. Brinkhurst &amp; Jamieson 1971, Brinkhurst 1986, Kathman &amp; Brinkhurst 1998). In a collection from this region (USNM 32980), reproductive organs are not visible (although one specimen has penial chaetae), and worms have very long hair chaetae in II (up to 1200 µm long, 1.8–2 times longer than the following ones), in contrast to the R. montana type series.</p><p>Anterior pectinates of R. longichaeta have only slightly longer distal teeth, in contrast to the distinctly longer and thicker distal tooth in the type material of R. montana (Fig. 16 A, D–E). Nevertheless, this character is usually difficult to interpret due to orientation of the pectinate chaetae, which are variably rotated. Immature rhyacodrilines with long hairs in segment II studied in other western localities (see other material above) are now provisionally ascribed to R. longichaeta . Anterior pectinates in these populations are variable; in some specimens the distal tooth is clearly longer than the lower, but not in others. Hispid hair chaetae are seen in some populations at 400x, but in individuals from Guadalupe Creek 1000x magnification is required, even with Nomarski microscopy. Hispid hair chaetae are rare in the genus, although they have been also described in R. hiemalis Ohtaka, 1995 .</p><p>Rhyacodrilus sodalis (Eisen, 1879) (Figs 17, 18)</p><p>Ilyodrilus sodalis Eisen, 1879: 11 . Eisen 1885: 887, Pl. VI, Fig. 5.</p><p>Rhyacodrilus sodalis (Eisen): Brinkhurst 1963: 53, 59.</p><p>Non Rhyacodrilus sodalis (Eisen): Brinkhurst 1965: 144, Fig. 8 E–K; Ohtaka 1995: Fig. 8 B, C).</p><p>Neotype. USNM 1202075, dissected worm stained in Harris' hematoxylin and slide-mounted in Canada balsam.</p><p>Paraneotypes. USNM 1202076-77, 1 dissected worm stained in Harris' hematoxylin, 1 sagittally sectioned and stained with hematoxylin and eosin.</p><p>Further material, not included in description. Former neotype (USNM 32644) and 4 paraneotypes (USNM 32645 No.0004, 4 slides) of R. sodalis sensu Brinkhurst (1965) from Lake Tahoe, invalidated (see below).</p><p>Locality. Tributary to Mountain Lake, Camino del Mar, N37.7879° W122.4698°, San Francisco, California, USA, Eisen's original type locality of the species (31 May 2011).</p><p>Description. Number of segments 58–62. Body diameter about 0.55 mm at VIII, to 0.7 mm at clitellum.</p><p>Prostomium rounded. Epithelium in anterior segments up to 18 µm high. Clitellum annular from about 1/ 2 X to XII, absent only around male pores, up to 52 µm high, formed by small glandular cells (Fig. 17 A). Male pores in posterior part of segment XI, within a shallow, lenticular depression; spermathecal pores at the very beginning of segment X, close to intersegment 9/10; both male and spermathecal pores about in line with ventral chaetae. One pair female pores, lateral to the line of ventral chaetae. Anchorage bridge inconspicuous or absent, at most a small ridge of slightly thickened epidermis between spermathecal pores.</p><p>Dorsal bundles with hair and pectinate chaetae: 2–4 hairs in anterior bundles (the longest ones 190–410 µm), 1–2 thin hairs back to segment XVI–XXV, hairs absent posteriorly; 4–5 pectinate chaetae (83–132 µm) per bundle, with divergent lateral teeth of equal length and 4–5 intermediate teeth in anterior segments; 2–3 sigmoid pectinates in postclitellar segments (79–87 µm) with 3–4 short intermediate teeth, distal teeth shorter than proximal (Fig. 18 D, E), nodulus at 1/3 from the tip. Ventral bundles with 4–6(7) bifid chaetae in anterior segments (94–132 µm), 2– 5 in postclitelar segments (81–90 µm); teeth equally long or with distal tooth shorter and thinner than proximal; in most ventral bundles some chaetae have short thin intermediate teeth, the most posterior segments with only 1 bifid chaeta per bundle (Fig. 18 F–H). Median to the male pore a bundle of 3–4 penial chaetae (90–120 µm long, the longest in the middle of the bundle), bifid with short teeth, nodulus at 1/3 from the tip, arranged fanwise (Fig. 18 I, J).</p><p>Dorsal pharyngeal pad well-developed. Pharyngeal glands from segment IV to anterior part of segment VII. Numerous granulated and nucleated coelomocytes in body cavity (diameter 12–24 µm) (Fig. 18 A). One pair testes in segment X and one pair ovaries in segment XI. Sperm sacs extend forward into IX and back to segment XVIII– XX. Ovisac back to XIX–XXII. Paired ventral glands observed in segments V to X, best developed in segment IX, located behind ventral chaetae (Fig. 18 B).</p><p>One pair spermathecae, composed of a short bulbous duct (72–85 µm long, 52–72 µm maximum diameter) with columnar epithelium, and an irregular ampulla filled with bundles of sperm, characteristically folded (as in the original description of the species) around a strong muscular strand attached to the duct and to the dorsal body wall (Fig. 17 B: ms; 17C). One pair male ducts in segment XI. Sperm funnel on septum 10/11, followed by a long (to 700 μm), convoluted vas deferens, diameter 23–38 µm (thinner at both ends), apically joining the atrial ampulla (Fig. 17 B). Atrium with long duct (95–150 µm long, 27–30 µm wide), associated with several muscular bundles attached to the ventral body wall, and an elongated, curved ampulla (ca. 145–270 µm long, 70–125 µm maximum diameter). The ampulla may appear globular if compressed in whole mounted specimens, or elongated and curved (originally described as "crescent shape" [Eisen 1885: 888]) if relaxed or after removing from the segment through dissection (Fig. 17 B, D). As reported by Eisen (1885: 888), the atrium is "extremely pellucid" (transparent): atrial musculature very thin (ca. 1–3 µm), atrial epithelium thick (15–40 µm high), granular, and faint, although distinct in histological sections, with small, peripheral nuclei (Fig. 17 E). Ampulla covered by layer of prostate cells, 50–70 µm high, forming numerous clusters, joining atrium individually. In histological sections, a very short, narrow penis (?) was observed within the lumen of the atrial duct (Fig. 17 F). Anterior ends of spermatozoa in male duct and spermathecal ampulla have a distinctly wavy appearance (corkscrew shaped?) (Fig. 18 C).</p><p>Remarks. The species was described by Eisen (1879, 1885) as Ilyodrilus sodalis, from a small tributary to Mountain Lake in San Francisco (California). The species was transferred to the genus Rhyacodrilus by Brinkhurst (1963: 53, 59) based on atrial morphology, although regarded as a species dubia. Eisen's types collection disappeared after the earthquake in 1906 and a neotype was designated and described by Brinkhurst (1965). A sampling by Brinkhurst at the type locality was not successful in getting more material of the species (R.O. Brinkhurst, pers. comm. 2011), and the neotype was instead selected from Lake Tahoe. The new type series was however of limited value since neither male ducts nor spermathecae were observable (see Brinkhurst 1965: 145). Therefore, although the taxonomic status of the species has been considered uncertain (Brinkhurst 1986), this has not been an impediment to including the species in several guides, using keys based on chaetal characteristics. As a result, R. sodalis has been widely reported in North America (Kathman &amp; Brinkhurst 1998).</p><p>Examination of Brinkhurst's neotype (USNM 32644) and paraneotypes from Lake Tahoe (USNM 32645 No.0004, 4 slides) did not provide any information on the structure of reproductive organs, probably due to the mounting media (most likely lactophenol). In the neotype, anterior pectinates have very long, thin teeth (Fig. 18 K, L) and there are 2 single-pointed penial chaetae, arranged in parallel. Lake Tahoe is a large, deep (499 m depth) water body, located at 1,897 m a.s.l. with highly oligotrophic waters (Gardner et al. 2000), while Eisen's original locality is in a very small, low elevation coastal seep, currently filled with vascular plants and draining an urbanized watershed. The linear distance between these two sites is about 230 km. The International Code of Zoological Nomenclature (ICZN 1999) states in Art. 75.3.6. that the paper in which a neotype is designated must state that "the neotype came as nearly as practicable from the original type locality and, where relevant, from the same geological horizon or host species as the original name-bearing type ". The poor condition of the Tahoe specimens, the lack of definition of the reproductive organs, and the fact that Brinkhurst's neotype series came from a water body very different in ecological and geological characteristics from the original indicates that the Lake Tahoe neotypes are invalid. Therefore we designate a new R. sodalis neotype from Eisen's type locality, adding details of morphological characters currently used in rhyacodriline descriptions. This description validates most of the morphological features formerly described by Eisen from the same locality.</p><p>The original description of R. sodalis gives little information on the shape of chaetae; however, the striated inner surface of the so-called "pseudo-comb-like spines" can be interpreted as the presence of intermediate teeth in dorsal chaetae, probably not clearly visible through Eisen's microcopy. Dorsal crochets are V-shaped with equal teeth in the San Francisco population (both in Eisen's description [1885: Fig. 5 e,f] and in the present paper), instead of the long, thin teeth seen in the Lake Tahoe population. Large and abundant coelomocytes, a character that is diagnostic of the subfamily Rhyacodrilinae (Hrabĕ 1963b), were not originally described by Eisen; they are present in the new neotype series, but this was likely not considered to be a diagnostic character at the time of the original description. There is a general agreement between Eisen's and the present descriptions of the spermatheca and male duct: the spermathecae have a short, bulbous duct, similar to many other Rhyacodrilus species, and an ampulla which appears twisted or folded; in the new material the ampulla is folded around a strong, dorso-ventral muscular strand originating at the spermathecal duct (Fig. 17 B, C). The male duct has a long, coiled vas deferens that apically joins the elongated atrium, and the long atrial duct opens in a simple pore. The conical structure that surrounds the atrial duct in the original description (incorrectly named oviduct by Eisen) is interpreted here as the covering of the atrial duct by numerous muscle strands attached to the ventral side of the body. The "pellucid" appearance of copulatory organs reported by Eisen is here interpreted as a vague or almost transparent atrial epithelium and very thin musculature. The most problematic issue is the lack of mention of modified penial chaetae in the original description of the species, a rather obvious character that is present in the specimens recently found in the type locality.</p><p>Based on the shape of the chaetae, presence of multiple penial chaetae but not spermathecal chaetae, the long vas deferens, and the atrial duct shorter than the ampulla, R. sodalis is most closely related to R. coccineus and a complex of similar species (see the key in this paper), some of which have been put in synonymy with the latter (e.g. Timm 1997). Rhyacodrilus sodalis has also been considered a junior synonym of other Rhyacodrilus species in this group. The synonymy of R. sodalis with R. sinicus Chen (e.g. Brinkhurst 1965, Brinkhurst &amp; Jamieson 1971, Qi &amp; Erséus 1985) is difficult to accept. In the original description by Chen (1940), the atrium is ovoid and the duct opens in a tegumentary fold, suggesting a closer relationship to R. coccineus . Rhyacodrilus sodalis has also been regarded as a possible junior synomym of R. coccineus (e.g. Timm 1997). Any proposal for synonymy of a species with the widespread R. coccineus requires a previous reconsideration of the taxonomic status of the latter, since at present the morphology of the species shows too much variability among the numerous descriptions by different authors in many parts of the world. At this point, we distinguish R. sodalis from R. coccineus mainly by the relatively long vas deferens joining the atrium apically, and by the more elongate, curved atrial ampulla described by Eisen and confirmed in the new material.</p></div>	https://treatment.plazi.org/id/154FD01D4B48FFF7FF6E9116BB80CF03	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodriguez, Pilar;Fend, Steven V.	Rodriguez, Pilar, Fend, Steven V. (2013): New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1-44, DOI: 10.11646/zootaxa.3664.1.1
