Leodamas Kinberg 1866
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 sing...
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| Summary: | 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 whe : Published as part of Blake, James A., 2017, Polychaeta Orbiniidae from Antarctica, the Southern Ocean, the Abyssal Pacific Ocean, and off South America, pp. 1-145 in Zootaxa 4218 (1) on pages 48-59, DOI: 10.5281/zenodo.245827 : {"references": ["Kinberg, J. (1866) Annulata nova. Ofversight af Kungliga Vetenskaps - Adakemiens Forhandlingar, Stockholm, 22, 239 - 258.", "Chamberlin, R. V. (1919) The Annelida Polychaeta. Memoirs of the Museum of Comparative Zoology, Harvard, 48, 1 - 514, plates 1 - 80.", "Hartman, O. (1957) Orbiniidae, Apistobranchidae, Paraonidae and Longosomidae. 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