Genus Leodamas Kinberg, 1866

Type-species: Leodamas verax Kinberg, 1866, by monotypy.

Synonym: Branchethus Chamberlin, 1919. Type-species: Branchethus latum Chamberlin, 1919, by monotypy. Fide Hartman 1957.

Diagnosis. Prostomium pointed on anterior margin, usually prolonged; most species with a single achaetous peristomial segment; immature adults of some species with two achaetous peristomial segments and adults of at least one species with vague indication of two achaetous segments. Branchiae single or multiple branches, either from anterior thoracic setigers 4–7 or from posterior thoracic setigers or first abdominal setigers. Posterior thoracic setigers with 0–2 postsetal lobes and 0–2 subpodial lobes, never more than four lobes of both types combined; not forming ventral fringes. Thoracic neuropodial uncini large, conspicuous arranged in one to many distinct vertical rows, with accompanying capillaries few or entirely lacking; heavy spear-like spines and bristle-topped setae absent. Abdominal neuropodia with projecting aciculae, either thin and inconspicuous or large and curved apically. Abdominal noto- or neuropodial flail setae present or absent.

Etymology. The generic name Leodamas Kinberg, 1866, is formed from Leo, Greek for lion, and dama, Latin for deer. It seems likely that Kinberg noticed the branches of the thoracic notopodial lamellae and compared them with antlers of a deer, hence the name. Leo is masculine, dama can be both masculine and feminine, but I believe this genus name is masculine as evidenced by the masculine name of the type species, verax.

Remarks. Leodamas was raised to full generic status by Blake (2000). A phylogenetic analysis demonstrated that Leodamas species were more closely related to species of Orbinia, Phylo, and Naineris than to Scoloplos sensu stricto. Previously, Leodamas was regarded as a subgenus of Scoloplos. The chief difference between species of Scoloplos and Leodamas is with the form and arrangement of the thoracic neuropodial uncini. In species of Scoloplos the thoracic neuropodial uncini are relatively thin, inconspicuous, and accompanied in each setal row by numerous capillaries. In contrast, the thoracic neuropodial uncini of all species of Leodamas are large, conspicuous and arranged in 1–7 (usually 1–4) distinct vertical rows; capillaries if present, are few in number, in separate fascicles, usually dorsal to the uncini, and relatively inconspicuous. One unique feature of several species of Leodamas, including the type-species, L. verax, is that the anterior row of uncini often extends ventrally, curving under other rows of uncini, continuing dorsally as a short posterior row. Other species of Leodamas confirmed to have this feature include L. cirratus, L. cochleatus, L. cylindrifer, L. hyphalos n. sp., L. maciolekae n. sp., L. marginatus, and L. tribulosus. By having such large and heavy thoracic neuropodial uncini, species of Leodamas are more similar to species of Naineris, Orbinia, and Phylo than to Scoloplos. Most species of Leodamas have emergent neuropodial aciculae in abdominal segments. The appearance of branchiae on anterior thoracic setigers in most of the better known species of Leodamas is usually considered as another way to separate these species from Scoloplos where branchiae begin more posteriorly. However, some Leodamas species with rows of heavy thoracic neuropodial uncini do have branchiae first present from a more posterior setiger.

In order to better understand the relationships of the South American and Antarctic species of Leodamas encountered as part of this study, species currently assigned to Leodamas either as a full genus or as a subgenus to Scoloplos were reviewed based largely on the literature. This review suggests that species of Leodamas may be divided into two groups (Table 1): (A) Species with branchiae beginning from an anterior thoracic setiger (4–7) and with the thoracic neuropodial uncini typically occurring in three or more vertical rows and (B) Species with branchiae beginning from a posterior thoracic or anterior abdominal setiger and with thoracic neuropodial uncini typically occurring in only 1 or 2 vertical rows. In general this dichotomy holds up well, however L. acutissimus (Hartmann-Schröder, 1991) is intermediate with branchiae beginning from a posterior thoracic segment and with 3–4 vertical rows of thoracic neuropodial uncini. The following list includes all known species of Leodamas either derived from the literature or encountered in this study. Table 1 includes the main morphological characters as taken from key references or from actual observations. There are likely additional species of Scoloplos that may eventually be reassigned to Leodamas. The following 29 species of Leodamas are currently recognized.

Species with branchiae from an anterior thoracic setiger (4–7); thoracic neuropodial uncini in 3–4 or more vertical rows:

Leodamas verax Kinberg, 1866). Type species

Leodamas brevithorax (Eibye-Jacobsen,2002) New combination Leodamas chevalieri (Fauvel, 1902) New combination Leodamas cochleatus (Ehlers, 1900) New status

Leodamas cirratus (Ehlers, 1897) New combination

[Aricia ohlini Ehlers, 1897] New synonymy

Leodamas dendrocirrus (Day, 1977) New combination Leodamas dubius (Tebble, 1955)

[Scoloplos (Leodamas) rubra australiensis Hartmann-Schröder, 1979] Fide Eibye-Jacobsen 2002 Leodamas fimbriatus (Hartman, 1957) New combination Leodamas gracilis (Pillai, 1961) New combination Leodamas hamatus Dean & Blake, 2015

Leodamas hyphalos n. sp.

Leodamas johnstonei (Day, 1934) New combination

[Scoloplos (Leodamas) uniramus Day, 1961] Fide Day 1977 Leodamas latum (Chamberlin, 1919) New combination Leodamas marginatus (Ehlers, 1897)

[Aricia marginata mcleani Benham, 1921] New synonymy

[Scoloplos (Leodamas) naumovi Averincev, 1982] New synonymy Leodamas orientalis (Gallardo, 1967) New status

Leodamas perissobranchiatus n. sp.

Leodamas rubrus (Webster, 1879)

Leodamas thalassae (Amoureux, 1982) New combination Leodamas tribulosus (Ehlers, 1897)

[S. armiger trioculata Hartmann-Schröder, 1962b] New synonymy)

Species with branchiae from a posterior thoracic setiger or anterior abdominal setiger (12–40); thoracic neuropodial uncini in 1–2 vertical rows.

Leodamas acutissimus (Hartmann-Schröder, 1991) New combination Leodamas agrestis (Nonato & Luna, 1970) New combination Leodamas cylindrifer (Ehlers, 1904) New combination

[Scoloplos (Leodamas) dendrobranchus (Hartman, 1957)] Leodamas maciolekae n. sp.

Leodamas madagascarensis (Fauvel, 1919) New combination Leodamas mazatlanensis (Fauchald, 1972) New combination Leodamas minutus López, Cladera & San Martín, 2003

Leodamas platythoracicus López, Cladera & San Martín, 2003 Leodamas texana (Maciolek & Holland, 1978) New combination Leodamas treadwelli (Eisig, 1914)

Incertae sedis Alcandra robustus Kinberg, 1866.

As part of this review several potential taxonomic problems have been identified, largely associated with the nature of the abdominal neuropodial uncini, including their size and the degree of curvature of the curved or hooked tip. Leodamas dubius was originally described from West Africa by Tebble (1955). However, the original account was brief and certain key characters were not clearly discussed or illustrated. The species was subsequently reported from Viet Nam (Gallardo 1967), the Andaman Sea (Eibye-Jacobsen 2002), and Australia (Hartmann- Schröder 1979 as L. rubra australiensis fide Eibye-Jacobsen 2002) and Zhadan et al. (2015). The two latter accounts reported considerable variability in the size, shape, and degree of curvature of the abdominal neuropodial uncini. It is likely that the original West African account and the more recent reports from Asia and Australia represent different species. The only resolution to this would be to re-examine the specimens reported by Tebble (1955).

Another potential problem involves L. johnstonei which was originally described from southern Africa by Day (1934) and has subsequently been reported from Australia (Day 1977). Here the main issue seems to be with the presence of 1 or 2 subpodial papillae in the last thoracic and anterior abdominal segments. Other aspects of the morphology have not been so carefully compared, and given the great geographic distance between records of the species it would be of interest to compare African and Australian specimens.

Scoloplos (Leodamas) mazatlanensis described by Fauchald (1972) from deep water off western Mexico is here referred to Leodamas, but the species is not well described and illustrated and may not belong to this genus. The arrangement of the thoracic neuropodial uncini is not stated, only that 10–15 uncini are present per neuropodium. Further, the text suggests that there are numerous capillaries in these neuropodia as well, which if true would imply that the species belongs in Scoloplos sensu stricto. The original specimens need to be reexamined.

Leodamas latum was described from off Panama in 588 m by Chamberlin (1919) as Branchethus latum and was later reported from off Burma in 457 m by Fauvel (1932) as Scoloplos latus. The species does not appear to have been reported since. There are differences in the two accounts. Neither author reported the number of rows of thoracic neuropodial uncini. The uncini are described by Chamberlin (1919) as strongly striated with numerous camerations, curved in a reverse direction and taper to a fine point. Fauvel (1932) on the other hand describes a more typical stout, blunt-tipped acicula that is relatively straight, and with transverse ribs on the convex side. Chamberlin (1919) did not report an emergent abdominal neuropodial acicula. Fauvel (1932) described a stout, blunt acicula accompanied by 4–6 capillaries. Given that the two accounts are so disjunct geographically and that morphological differences are reported, it is possible that more than one species is involved and the collections, if available should be re-examined. A closely related new species, L. perissobranchiatus n. sp. differs in that the branchiae begin on setiger 4 instead of 5 and the thoracic neuropodial uncini have a lateral sheath (see below).

Scoloplos (Leodamas) rubra orientalis Gallardo, 1967 is here elevated to full species status. Leodamas orientalis has a ventral cirrus on the abdominal neuropodia, whereas L. rubrus has an elongated postsetal lobe. This also represents another very disjunct distribution: US Atlantic and Gulf coasts for L. rubrus and SE Asia for L. orientalis.

Another interesting species pair is Leodamas gracilis described by Pillai (1961) from Sri Lanka, Viet Nam by Gallardo (1967), the Andaman Sea by Eibye-Jacobsen (2002); and Leodamas agrestis described by Nonato & Luna (1970) from off NE Brazil in 20–50 m. Both species have 3–4 large pointed acicular spines anterior to the smaller and normal vertical rows of uncini that occur in thoracic neuropodia. The two species differ however, in that the vertical rows of uncini number only one in L. agrestis and 3–4 in L. gracilis. In addition, the branchiae of L. agrestis begin on the first abdominal segment (setiger 16) whereas branchiae begin on setiger 6 in L. gracilis. Thus, although these are the only two species of Leodamas reported with large anterior acicular spines anterior to the vertical rows of uncini of thoracic neuropodia, L. gracilis is related to the species in Table 1 Group A and L. agrestis is in Group B.

Another species with an interesting history is Scoloplos cylindrifer originally described from South Island, New Zealand by Ehlers (1904). Ehlers had complete specimens up to 17 mm long and 115 setigers. This original description depicted a fairly typical orbiniid with a pointed prostomium, peristomium with a single ring, and parapodia consisting elongate, flattened notopodial lamellae and reduced neuropodia. All setae were described as thin hairy bristles (“dünner feilkerbiger Borsten”), undoubtedly referring to the camerated nature of most orbiniid capillaries. The presence or absence of thoracic neuropodial spines or uncini was not indicated. The branchiae were described and illustrated as single, but one was noted to have a protrusion, suggesting bifurcation. The second report of the species was by Augener (1914) from intertidal sands in SW Australia based on two specimens, one of which was missing the anterior end. A specimen from New Zealand was provided from the Bremer Museum for comparison, but it was not stated if this was one of the syntypes from Ehlers’ collection. The Australian specimen was larger, 42 mm long and with 210 segments. The nature of the parapodia and setae were not mentioned. Instead, a detailed description of the branchiae was provided; these were determined to have 2, 3, and 4 branches. The one figure (Augener 1914: Plate I, fig. 4) clearly shows dichotomous branching. Augener (1926) recorded the species from near Dunedin, South Island, New Zealand. He noted that the branchiae were from setiger 22 and had up to four branches; no information was provided on the thoracic neurosetae. The fourth report of the species was by Monro (1939) based on several specimens from Tasmania, Australia. One complete specimen was 30 mm long with 140 setigers. All setae were reported as camerated capillaries leading Monro to transfer the species from Scoloplos to Haploscoloplos. He noted that branchiae were branched with up to five filaments. Hartman (1957) referred to the species as Haploscoloplos cylindrifer and reviewed the reports and noted considerable variability within the species accounts, in particular that the branchiae began anywhere from setigers 17–50. None of these earlier reports made any mention of spines or uncini in the thoracic neuropodia; but projecting uncini were observed by Ehlers (1904) in the posterior neuropodia, a characteristic typical of many species of Leodamas. Furcate setae were not reported in any of these early accounts.

In the same paper where she reviewed the history of Haploscoloplos cylindrifer, Hartman (1957) described a new species, Scoloplos (Leodamas) dendrobranchus from various intertidal habitats in South Australia. This was another species with dendritically branched branchiae with up to six filaments reported from setiger 18, or the first abdominal setiger. In addition to capillary setae, Hartman reported that this species lacked notopodial furcate setae but had 8–14 large thoracic neuropodial uncini arranged in a distinct J-shaped row anterior to the capillaries; these uncini were reported as blunt tipped, and with no hood. She also reported the presence of projecting aciculae in posterior neuropodia. All of these characters agree with the genus Leodamas as defined in this study for species of Group B, where large thoracic neuropodial uncini are present in 1–2 vertical rows and branchiae begin in posterior thoracic or anterior abdominal segments.

Day (1975: 1977) reported on eight specimens from South Australia that he identified as Scoloplos cylindrifer that differed from earlier accounts in having instead of lacking a small group of curved, serrated hooks in addition to crenulate capillaries in thoracic neuropodia. He also had Monro’s specimens in the British Museum checked and they were reported to him as also having hooks in thoracic neuropodia. For this reason, Day (1975) synonymized Hartman’s (1957) S. dendrobranchus with S. cylindrifer. Day (1977) later identified numerous additional specimens of S. cylindrifer from most coasts of Australia and considered it to be one of the most common orbiniids he encountered. In the same paper, Day (1977) determined that because Monro (1993a) had established Haploscoloplos cylindrifer as the type species of his genus Haploscoloplos Monro, that a new genus was required because the type species had an anterior row short hooks in the thoracic neuropodia in addition to capillaries and thus belonged to the genus Scoloplos. In addition to the Australian material, he examined a specimen from near Christchurch, South Island, New Zealand, near the type locality of the species and thus confirmed the presence of thoracic neuropodial hooks in the species from both New Zealand and Australia. He therefore proposed a new genus, Leitoscoloplos to include those remaining species formerly assigned to Haploscoloplos (see above for earlier account of Leitoscoloplos). Day & Hutchings (1979) summarized the Australian and other records for the species.

Hartmann-Schröder (1981) as part of a series of papers on Australian polychaetes provided the first detailed description of the thoracic neurosetae of Scoloplos cylindrifer since Hartman (1957) (as S. (Leodamas) dendrobranchus). The thoracic neurosetae included both capillaries and uncini. The latter were arranged in a horseshoe-shaped double row with the anterior row extending from the top of the fascicle vertically, and then curving under and extending part way up the posterior border of the setal fascicle becoming a short second row (Hartmann-Schröder, 1981: Fig. 101). This is similar to what Hartman (1957) called a J-shaped row. The uncini have thick shafts, narrowing to a blunt tip; a lateral sheath is present together with transverse ribs on the concave side of the shaft. The uncini on the anterior row have less distinct ribs on the shaft than the posterior uncini (Hartmann-Schröder, 1981: Figs. 104–105). As in previous reports, furcate setae were not observed. Hartmann- Schröder (1981) also noted that flail setae were absent, branchiae were first present from setiger 18 with up to five branches, and observed abdominal neuropodia with two protruding aciculae (Hartmann-Schröder, 1981: Fig. 103).

These observations by six different investigators eventually provided details sufficient to categorize the species. Based on the definition provided in the present study, the orbiniid species “ cylindrifer ” belongs to the genus Leodamas Group B and is included as such in Table 1. Leodamas cylindrifer is one of several species in the genus where a vertical anterior row of neuropodial thoracic uncini curves under other setae and continues again dorsally as a short posterior row.

The status of Alcandra robustus described by Kinberg (1866) is uncertain. The only specimen, an anterior fragment from Brazil with the head and eight setigers, was examined by Hartman (1948, 1957). The specimen was reported as having a conical prostomium, branchiae from setiger 5, only capillaries in both noto- and neuropodia of setiger 1, and with uncini in palisaded rows from setiger 2. Uncini were illustrated by Hartman (1957) as curved with a blunt tip and with transverse rows of ribs on the convex side of the shaft. Due to the fragmented nature of the holotype, this species cannot be fully compared with other species or even confirmed as belonging to Leodamas.

In the present study, eight species, three new to science are reported from South American and Antarctic seas. The type species, Leodamas verax is redescribed from new material off Argentina and a neotype is designated. Type specimens of five of Ehlers’ species have been examined and redefined, resulting in one being designated a synonym and another being resurrected from synonymy.

Species/Character Shape of thorax in X- No. thoracic setigerous No. rows of thoracic Capillaries with Structure of thoracic neuropodial

section segments neuropodial uncini thoracic neuropodial uncini uncini

Group A: Species of Leodamas with branchiae from anterior thoracic setigers; thoracic neuropodial uncini typically in 3 or more vertical rows

Leodamas verax Kinberg, Dorsoventrally flattened 22–24 3 vertical rows & 1 short Absent Blunt–tipped with groove on convex posterior row curving ventral to side; with weakly developed transverse

Type species) rows 1–3 ribs along shaft

Leodamas brevithorax Narrow, becoming depressed 15–17 4 vertical rows with first Few in dorsal tuft Uncini curved with rounded tip and Eibye– Jacobsen, 2002) or flattened from setiger 4 curving under 2–-3 and shaft with 5–12 transverse ridges

forming row 4

Leodamas chevalieri Flattened, broad 20–27 4–5 vertical rows Present, long, becoming Uncini straight, narrowing at tip, blunt, Fauvel, 1902) more numerous with lateral hood

Leodamas cochleatus Rounded, only weakly 10–29 (size related) 3 vertical rows & 1 short Absent Uncini of anterior setigers with straight

Ehlers, 1900) flattened dorsally posterior row curving ventral to shaft, blunt tip and weak transverse rows 1–3 ribs; transitioning to uncini with curved expanded tips with subapical notch and prominent ribs on shaft

Leodamas cirratus (Ehlers, Dorsoventrally flattened 21–32 3 long vertical rows & 1 short Absent Long, tapering; blunt-tipped with long) posterior row curving under vertical notch on concave side; weakly rows 1–3 developed transverses ribs on shaft

Leodamas dendrocirris Dorsoventrally flattened. 17–18 4–5 vertical rows; arrangement Crenulate capillaries in Long, curved apically with blunt tip

Day, 1977) not stated. single posterior row and transverse rows of ribs on convex

side.

Leodamas dubius Dorsoventrally flattened. 15–19 or 21–23 4–5; last row short Few with posterior row Long, curved apically to blunt tip; shaft

Tebble, 1955) uncini with transverse ribs; Thai and

Australian specimens with thickened hood, imparting bidentate appearance.

Leodamas fimbriatus Not stated. 24–30 Up to 4 vertical rows of uncini; Absent Uncini of first row longest, thickest,

Hartman, 1957) arrangement not stated. and with sharply curved, smooth tips; uncini of following rows thinner, less curved

Leodamas gracilis (Pillai, Short, depressed or flattened. 13–17 1 anterior row of 3–4 heavy Crenulate capillaries in Two types: 3–4 anterior heavy, simple) spines; 2–3 vertical rows of posterior row of uncini spines; second curved spines with numerous hooded uncini; first hooded tip and transverse rows of ribs row curves under rows 2–3. on shaft

……continued on the next page Species/Character Shape of thorax in X- No. thoracic setigerous No. rows of thoracic Capillaries with Structure of thoracic neuropodial

section segments neuropodial uncini thoracic neuropodial uncini

uncini

Leodamas hamatus Dean & Dorsoventrally depressed, 13–21 2–4 vertical rows & partial 5th 2–7 thin capillaries in Two types: anterior 4–5 setigers with

Blake, 2015 but not flattened. row dorsal tuft straight shaft, blunt tip and thin hood on convex side; uncini of following setigers with larger thickened hood, imparting bidentate appearance

Leodamas hyphalos n. sp. Inflated first 3–4 setigers, 16–18 4 vertical rows; first row 2–3 capillaries in Shaft curved to straight, tapering to then depressed but not curving ventral to 2 and 3, then superior position to last rounded apex; shaft with irregular flattened merging with posterior row row of uncini transverse ribs

Leodamas johnstonei (Day, Flattened dorsally, rounded 18–24 3–5 vertical rows Few in posterior row Shaft curving at tip; weakly serrated,

1934) ventrally with lateral flange

Leodamas latum Dorsoventrally flattened ~20 Not stated, but “setae arranged Present Elongate, curved to narrow tip, shaft Chamberlin, 1919) in vertical series.” with numerous transverse rows.

Leodamas marginatus Dorsoventrally flattened 14–20 3 vertical rows & 1 short 0–few in tuft superior to Blunt-tipped with sub-terminal groove Ehlers, 1897) after first 3 setigers transverse row curving ventral 3 vertical rows of uncini or notch & low transverse crenulated

to rows 1–3 ribs on shaft (SEM) Leodamas orientalis Broadly oval 18–21 4–5 vertical rows, 5th row from 1–2 in uppermost Curved, blunt-tipped, with up to 9 Gallardo, 1967) New middle thoracic setigers, only location transverse ribs on concave side status ½ as long as 1–4.

Leodamas Broad, dorsoventrally 11–13 3–4 vertical rows of uncini; Long, thin silky Distally curved, notched, with lateral perissobranchiatus n. sp. flattened arrangement not stated. camerated setae mixed flange and transverse rows of ribs

with uncini along shaft

Leodamas rubrus (Webster, Broadly oval, dorsoventrally 23–28 3–5; arrangement not stated. Few in uppermost Curved apically, with blunt tip and 1879) depressed location transverse ridges on convex side of

shaft

Leodamas thalassae Dorsoventrally flattened 12–15 2 curved rows of large uncini. A row posterior to Thick, curved apically, with blunt tip Amoureux, 1982) uncini and numerous transverse ridges on

shaft

Leodamas tribulosus Dorsoventrally flattened 22–25 5–7 dense vertical rows of Crenulated capillaries in Bent in posterior direction; tip with 2 Ehlers, 1897) uncini; first row curves under 2 rows posterior to pointed teeth followed by long groove rest of uncini, merging with uncini on convex side flanked by lateral; shaft short posterior row with rows of low transverse ribs. ……continued on the next page

Species/Character Shape of thorax in X- No. thoracic setigerous No. rows of thoracic Capillaries with Structure of thoracic neuropodial

section segments neuropodial uncini thoracic neuropodial uncini uncini

Group B: Species of Leodamas with branchiae from posterior thoracic or anterior abdominal setigers; thoracic neuropodial uncini typically in 1–2 rows

Leodamas acutissimus Only slightly flattened 14–20 3–4 rows Present in single row Uncini of anterior setigers smooth,

Hartmann-Schröder, 1991) posterior to last row of curved with 4–5 transverse rows of uncini denticles; uncini of posterior setigers straight with lateral sheath and no rows of denticles

Leodamas agrestis Nonato Broad, dorsoventrally 15 1 row, setigers 1–8; additional Present as dorsal tuft, Single row of uncini, curved, tapering Luna, 1970 depressed anterior row of 2–3 large, middle and posterior to pointed tip; large anterior spines heavy spines setigers 7–15 thoracic setigers short, thick, tapering to narrow tip

Leodamas cylindrifer Dorsoventrally depressed 14–17 A single J-shaped row with 8– Present with ca. 12 Simple, blunt, distally curved, with

Ehlers, 1904) 14 uncini curving ventrally capillaries in a vertical lateral sheath or hood; shaft with

under group of capillaries; fascicle transverse ribs uncini reduced to ~ 7 in

posterior thoracic setigers.

maciolekae n. sp. Flattened 40 2 rows, with 1st row curving Absent Simple, weakly curved, without

under 2nd. transverse ribs

madagascarensis Fauvel, Dorsoventrally flattened 24–30 2 rows, with 1st curving under Few in a dorsal tuft Simple, blunt, straight; without

2nd. transverse ribs

mazatlanensis (Fauchald, Weakly dorsoventrally 15 Not stated, but 10–15 uncini Present “numerous” Simple, curved, with 10–15 transverse) flattened present. shallow ridges on convex side. minutus Lopez, Cladera cylindrical 13 1–2 rows (set 1–3), 2 rows (set Absent setigers 1–9; Simple, straight with weak transverse San Martin, 2003 4–13) present from setiger 10 ribs

platythoracicus Lopez, Cylindrical anteriorly; flatted 19 1 (set 1) 1–2 uppermost in Simple, straight, without transverse ribs

Cladera & San Martin, 2003 posteriorly 2 (set 2–13) fascicle

1 (14–19)

texana (Maciolek & cylindrical 11–20 1 row of 5–9 uncini 1–few, in dorsal tuft Simple, weakly curved, rounded tip,

Holland, 1978)—Includes without transverse ribs

Naineris sp. A of Taylor

1984).

treadwelli (Eisig, 1914) flattened 14–22 2 rows in anterior and middle Absent Simple, weakly curved, rounded tip,

setigers; thereafter 1 row without transverse ribs ……continued on the next page Species/Character Emergent abdominal Abdominal Noto- Setiger Unique characters Geographic References noto- & neuropodial & neuropodial Branchiae Distribution

aciculae flail setae Begin

Leodamas verax Kinberg, Noto: Absent Absent 5–6 Thoracic notopodia 2–6 with single Off Uruguay and Hartman 1948b; 1957;

1866 Neuro: 1–2 projecting lobe; then with 2–4 separate branches Argentina, intertidal to This study

Type species) aciculae 62 m.

Leodamas brevithorax Noto: 2–3 thin, slightly Absent 6 Narrow thorax; low number of thoracic Andaman Sea, 17–79 m Eibye-Jacobsen 2002 Eibye– Jacobsen, 2002) emergent aciculae; setigers; thin emergent acicula in

Neuro: with 1 thin abdominal noto– and neuropodia

acicula

Leodamas chevalieri Noto: absent; Not stated 6 Furcate setae absent West Africa; Red Sea, Fauvel 1902, 1953; Fauvel, 1902) Neuro: 1 simple acicula Gulf of Aden, Arabian Gravier 1906; Wesenberg-

Gulf; Indian Ocean. Lund 1949

Leodamas cochleatus Noto: Absent Absent 6 Change in morphology of thoracic Offshore Argentina, 454 Ehlers 1900, 1901; This Ehlers, 1900) Neuro: 1–2 simple uncini from anterior spines with m; Chile, Straits of study

aciculae straight shaft, blunt tip and weak Magellan; 46 m.

transverse ribs to spines with curved

and expanded tips with subapical notch

and prominent ribs on shaft

Leodamas cirratus Noto: absent; Neuro: Absent 6 Single subpodial lobe on posterior SE Argentina; Falkland Ehlers 1897; Hartman Ehlers, 1897) with a single acicular thoracic and anterior abdominal Islands; Straits of 1957; This study

spine with or without thin segments; no capillaries accompany Magellan; South Orkney

hyaline hood thoracic neuropodial uncini. Islands; shallow

subtidal, 598 m. Leodamas dendrocirris Noto: Absent; Absent 5 Thoracic notopodia divided into 2–5 Australia, New South Day 1977 Day, 1977) Neuro: 1 long acicula separate branches continuing on Wales, 65 m.

with smooth bent tip. abdominal segments; thoracic

neuropodia with a single postsetal

lamella.

Leodamas dubius Noto: absent; Neuro, 1 Not stated 6–7 Furcate setae present; abdominal West Africa, 3–11 m; Tebble 1955; Eibye- Tebble, 1955) strongly hooked acicula. neuropodial uncini strongly hooked. Thailand; Australia, Jacobsen 2002; Zhadan et

Queensland. al. 2015

Leodamas fimbriatus Noto: a single acicula, Absent 7 3–4 subpodial lobes on posterior South Australia in sand; Hartman 1957; Day 1977 Hartman, 1957) slightly projecting; thoracic and anterior abdominal intertidal

Neuro: single aciculum setigers; these decreasing over about

projecting, slightly 15 abdominal segments, then

curved to blunt tip. disappearing.

……continued on the next page Species/Character Emergent abdominal Abdominal Noto- Setiger Unique characters Geographic References noto- & neuropodial & neuropodial Branchiae Distribution aciculae flail setae Begin

Leodamas gracilis (Pillai, Noto: absent; Absent 6 Presence of two kinds of thoracic Ceylon; Viet Nam, Pillai 1961; Gallardo

) Neuro: 1–2 simple, neuropodial uncini: 3–4 heavy, simple shallow water; Andaman 1967;

straight aciculae similar anterior spines; plus more numerous Sea, 19– 38 m. Eibye-Jacobsen 2002 to heavy spines of uncini in 2–3 vertical rows with

thoracic neuropodia hooded tip and transverse rows of ribs

on shaft.

Leodamas hamatus Dean Noto: absent; Absent 6 Heavy, curved uncini in abdominal Off Pacific Costa Rica, Dean & Blake 2015

Blake, 2015 Neuro: large, projecting neuropodia; change in structure of 11– 26 m.

acicula with hooked tip. thoracic neuropodial uncini from

anterior to posterior of thorax.

Leodamas hyphalos n. Noto: absent; Noto: 2 flail setae 6 Intersegmental annulations from Drake Passage in deep This study. Neuro: 1–2 blunt-tipped present; setigers 8–9; notopodial flail setae; water; 2888–4008 m. aciculae. Neuro: Absent

Leodamas johnstonei Noto: absent; Noto: absent; 6 Prostomium sharply pointed; 1–2 SW and South Africa; Day 1934, 1967, 1977;

Day, 1934) Neuro: 1 acicula, curved Neuro: 3–4 subpodial lobes in last thoracic Mozambique; Hartman 1957 at tip crenulate with fine segments and anterior abdominal intertidal.— Australia, hairlike tips setigers. Victoria, NSW, and Qld.

Leodamas latum Noto: absent Not stated. 5, stated as Anterior thoracic branchiae single, Off Panama, 588 m; off Chamberlin 1919; Fauvel

Chamberlin, 1919) Neuro: 1 blunt acicula somite 6 by then divided into 3–9 palmately Burma, 457 m. 1932; Hartman 1957

(Fauvel 1932) Chamberlin arranged branches through middle

abdominal segments; then number

decreasing again in posterior

abdominal segments

Leodamas marginatus Noto: absent; Absent 6 Neuropodial subpodial lobes absent. Southern South Ehlers 1897;

Ehlers, 1897) Neuro: 1 projecting America; widespread in Hartman 1957, 1966; acicula, distally curved sub-Antarctic and Knox 1998; This study Antarctic seas; intertidal to 1674 m

Leodamas orientalis Noto: absent Neuro: 1 Absent 6 Furcate setae with nearly equal tynes; Viet Nam; Andaman Gallardo 1967; Eibye-

Gallardo, 1967) New distally hooked acicula abdominal neuropodia with ventral Sea, 21– 79 m. Jacobsen 2002

status cirrus

Leodamas Noto: absent Absent 4 Branchiae single on anterior and Western Chile, 192 m. This study

perissobranchiatus n. sp. Neuro: tip of acicula middle thoracic segments; becoming

emergent palmately branched with 2, 3, and 4

branches in abdominal segments

Leodamas rubrus Noto: absent; Absent 6 Furcate setae with nearly equal tynes; E and SE United States; Webster 1879: Hartman

Webster, 1879) Neuro: 1 heavy spine Last thoracic and anterior abdominal Gulf of Mexico; 1951, 1957; Taylor 1984

with curved, smooth tip segments with long, tapering postsetal intertidal to 200 m. lobe.

Leodamas thalassae Noto: 1 present, pointed; Not stated 5 Prominent postsetal lobe in thoracic NE Atlantic, off Brittany Amoureux 1982

Amoureux, 1982) Neuro: 1 present, pointed neuropodia; relatively few thoracic coast of France, 850– neuropodial uncini. 1400 m.

Leodamas tribulosus Noto: absent; Absent 5 Large number of thoracic uncini each Western South America, Ehlers 1897; This study

Ehlers, 1897) Neuro: 1–2 aciculae with curved posteriorly and with dorsal Patagonia; Argentina; blunt tips. groove terminating in two pointed intertidal to shallow teeth. subtidal

Leodamas acutissimus Noto: not emergent; Noto: absent; 16–18, Thoracic neuropodial uncini transition Australia, Gladstone Hartmann-Schröder, 1991;

Hartmann-Schröder, Neuro: thin curved Neuro: present posterior from 3–4 rows of smooth curved and Lizard Island, Zhadan (2015)

1991) spines, with tips thorax spines with transverse ribs to straight Qld,

emergent spines with lateral sheath and no ribs.

Leodamas agrestis Noto: not emergent; Absent 16, 1st Last 9–10 thoracic setigers with three Off Brazil, 2–100 m Nonato & Luna 1970

Nonato & Luna, 1970 Neuro: not emergent abdominal large bent, pointed, acicular spines

anterior to single row of pointed uncini

Leodamas cylindrifer Noto: imbedded; Neuro: Absent 18, Dendrically branched branchiae with New Zealand; Ehlers 1904; Augener 1914;

Ehlers, 1904) long, sharply curved on abdominal up to 6 filaments. Thoracic Australia, intertidal Hartman 1957; Day 1977; tip neuropodial uncini in a single anterior Hartmann-Schröder 1981 row curving ventrally under row of

capillaries.

maciolekae n. sp. Noto: single spine; Noto: Absent; 29, thoracic Large number of thoracic setigers (40); Argentina, 14 m This paper Neuro: single thin Neuro: Present, with abdominal neuropodial flail setae

straight acicula with tapering

mucron-like tip

madagascarensis Noto: not emergent; Absent 22, thoracic Large number of thoracic setigers (24– SE Africa, Fauvel 1919; Day 1951, 1967

Fauvel, 1919 Neuro: long acicula, tip 30); thoracic neuropodial uncini simple Madagascar emergent straight, with few capillaries

mazatlanensis Noto: not emergent; Not stated 12 Not well described; thoracic Off Western Mexico, Fauchald 1972

Fauchald, 1972) Neuro: 3–4 curved, neuropodial with fingerlike postsetal 2487–2560 m project lobe; abdominal neuropodia with

ventral cirrus.

minutus Lopez, Noto: not emergent; Absent 12–16 Cylindrical thorax, not flattened; Pacific Panama, Lopez et al. 2003

Cladera & San Martin, Neuro: sigmoid acicula setigers 1–3 uniramous, notosetae intertidal

absent.

platythoracicus Lopez, Noto: not emergent; Absent 20, 1st Thorax cylindrical setigers 1–5, then Pacific Panama, Lopez et al. 2003

Cladera & San Martin, Neuro: single, brown, abdominal distinctly flattened; thoracic neuro intertidal

straight acicula uncini in 1 row on setiger 1, then 2

rows, again reduced to 1 row setigers

14–19.

texana (Maciolek & Noto: not emergent; Absent 2nd to 4th Thoracic neuropodial uncini in single NE Colombia; Gulf Maciolek & Holland 1978;

Holland, 1978)— Neuro: narrow, thin, tip abdominal row throughout of Mexico, Taylor 1984; Granados-

Includes Naineris sp. A emergent Louisiana-Texas, 2–5 Barba & Solís-Weiss 1997b

Taylor (1984). m

treadwelli (Eisig, Noto: not emergent; Absent Last Thoracic neuro uncini transition from 2 Caribbean; Puerto Treadwell 1901; Hartman

) Neuro: not emergent, or thoracic or rows to 1 row anterior to posterior. Rico; NE Colombia; 1957; Maciolek & Holland only the tip 1st Pacific Mexico, 6– 1978; Leon-González & abdominal 220 m Rodríquez 1996; Granados- Barba & Solís-Weiss 1997a

Accounts from West Africa (Tebble 1955) and Indo-Pacific (Eibye-Jacobsen 2002; Zhadan et al. 2015) likely represent different species. (2)Accounts from Panama (Chamberlin 1919) and

Burma (Fauvel 1932) may represent different species.