A Review of the Genus Eunice - Smithsonian Institution Libraries
A Review of the Genus Eunice - Smithsonian Institution Libraries
A Review of the Genus Eunice - Smithsonian Institution Libraries
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NUMBER 523 15<br />
be longer than <strong>the</strong> o<strong>the</strong>r teeth (compare Figures 13a, 5h, and<br />
9c). All teeth may be wide and abruptly tapering distally<br />
(Figure 21c) or evenly tapering from <strong>the</strong> bases (Figure 5h). The<br />
number <strong>of</strong> pectinate setae usually increases from anterior to<br />
posterior setigers; in some species, pectinate setae appear to be<br />
absent in <strong>the</strong> first 15-20 setigers.<br />
Compound falcigers are usually distinctly narrower than<br />
aciculae or subacicular hooks, but in some species <strong>the</strong>y are as<br />
thick as <strong>the</strong> latter. Falcigers are composed <strong>of</strong> shafts and<br />
appendages. The shaft is usually slender, but <strong>of</strong>ten expanded<br />
distally to a distinct head (compare Figures 23a and 4e). Shaft<br />
heads may be marginally dentate or serrated. Shafts are usually<br />
clear, nearly translucent, but may become more strongly<br />
sclerotinized, especially towards <strong>the</strong> bases, which <strong>the</strong>n become<br />
chestnut, brown, or even black in color (e.g., <strong>Eunice</strong> sebastiani).<br />
The appendages may be short (about as long as <strong>the</strong><br />
distally expanded head <strong>of</strong> <strong>the</strong> shafts, Figure 4e) or <strong>the</strong>y may be<br />
distinctly longer, becoming in some instances, very long and<br />
narrow (Figure 8b). The appendages may distinctly taper<br />
towards <strong>the</strong> tip (Figure 8b) or have nearly to completely parallel<br />
sides (Figure 12f). Distally, <strong>the</strong>y arc usually bi- or, rarely,<br />
tridentate. In bi- and tridentate falcigers (Figures 4e and 5e), <strong>the</strong><br />
teeth are in a row, with one above <strong>the</strong> o<strong>the</strong>r. The teeth may be<br />
nearly erect or variably curved and differ in size from barely<br />
noticeable to long and pointed. Reduction in size <strong>of</strong> <strong>the</strong> teeth is<br />
most common in <strong>the</strong> proximal teeth. Distally <strong>the</strong> appendages<br />
are covered by a pair <strong>of</strong> guards. The guards usually fit closely<br />
around <strong>the</strong> appendages, but are very much wider in a few<br />
species. Each guard consists <strong>of</strong> a convex membrane, which<br />
distally may be rounded or bluntly to sharply pointed (Figures<br />
3b, 4e, 5e). The guards may be equipped with distinct mucros<br />
(Figure 130- Such mucros are mostly present in forms with<br />
bluntly to sharply pointed guards, but may also be present in<br />
forms with rounded guards (Figure 16f,i). The edge facing<br />
away from <strong>the</strong> teeth may be slightly thickened and <strong>the</strong> o<strong>the</strong>r,<br />
cutting edge may be serrated.<br />
Pseudocompound falcigers (Figure 7h, j,k; Figure 86h,i) have<br />
thick shafts that are barely inflated near <strong>the</strong> reduced joint. The<br />
joint may be recognizable only as a swelling on one side <strong>of</strong> <strong>the</strong><br />
falciger or be more distinct. The pseudoappendages are short,<br />
with nearly parallel sides and bidentate. Both teeth are directed<br />
distally and tapering.<br />
Shafts <strong>of</strong> <strong>the</strong> compound spinigers closely resemble those <strong>of</strong><br />
<strong>the</strong> falcigers (Figure 57c). Appendages (blades) vary in length<br />
and are always knife-edged; <strong>the</strong> narrower edge may be serrated.<br />
Each neuropodium may be supported by a single acicula;<br />
perhaps most common are paired aciculae, but up to about<br />
seven have been recorded (e.g., <strong>Eunice</strong> manihine, four in Figure<br />
69e). When multiple aciculae are present, <strong>the</strong>y are arranged in<br />
a dorsoventral row. The aciculae usually are thicker than all<br />
o<strong>the</strong>r setae. The number and shape may vary from anterior to<br />
posterior end. Multiple aciculae are more common in anterior<br />
than in posterior setigers. Most species have tapering, distally<br />
bluntly pointed, straight aciculae; some species have distally<br />
hammer-headed or irregularly thickened and truncated aciculae.<br />
In some species, aciculae are distally bifid. Gently curved,<br />
or geniculate aciculae are common. Aciculae from a single<br />
neuropodium may differ in shape.<br />
Subacicular hooks are exclusively present in <strong>the</strong> neuropodia<br />
and are first present from setigers 15-35 in most species.<br />
Hooks are usually present in all remaining setigers. Hooks<br />
originate near <strong>the</strong> base <strong>of</strong> <strong>the</strong> aciculae, but form a distinct angle<br />
relative to <strong>the</strong> aciculae (Figure 3e) and emerge along <strong>the</strong> lower<br />
edge <strong>of</strong> <strong>the</strong> neuropodia. Subacicular hooks are usually slightly<br />
thinner than <strong>the</strong> aciculae and are <strong>of</strong>ten somewhat lighter in<br />
color. Most species have a single subacicular hook per<br />
parapodium, but in some species a vertical series <strong>of</strong> as many as<br />
5 may be present (e.g., <strong>Eunice</strong> vittata, three in Figure 114e).<br />
Distally, subacicular hooks may be bidentate (Figures 3d and<br />
7c,h), tridentate (Figures 4d, 5c,d) or simple and spine-like<br />
(Figure 103k). In some species <strong>the</strong> two distal teeth in <strong>the</strong><br />
tridentate hooks are arranged in tandem (e.g., E. cirrobranchiata,<br />
Figure 31e; E.flaccida, Figure 46c), but in most cases<br />
<strong>the</strong> teeth are crested (i.e., with <strong>the</strong> tips <strong>of</strong> <strong>the</strong> teeth in a single<br />
plane parallel to <strong>the</strong> curvature <strong>of</strong> <strong>the</strong> setal axis, e.g., Figures 4d,<br />
5c,d).<br />
Structurally, aciculae and subacicular hooks are similar<br />
when viewed with a light microscope; when any internal<br />
structure is visible, <strong>the</strong>y appear to consist <strong>of</strong> an inner core <strong>of</strong><br />
dense fibers, covered by a non-fibrous sheath. In some species,<br />
<strong>the</strong> separation between core and sheath is distinct, with a<br />
dark-colored core and a clear, <strong>of</strong>ten translucent sheath. In o<strong>the</strong>r<br />
species <strong>the</strong> separation between core and sheath is indistinct.<br />
Aciculae and subacicular hooks may be clear and nearly<br />
colorless, yellow, chestnut colored, various shades <strong>of</strong> brown,<br />
and in some instances a very distinct jet black. If paired<br />
aciculae are present, <strong>the</strong>y are <strong>of</strong>ten <strong>of</strong> different colors. If paired<br />
subacicular hooks are present, <strong>the</strong>y usually have <strong>the</strong> same<br />
color, but replacement subacicular hooks that have yet to<br />
emerge are <strong>of</strong>ten lighter in color. Aciculae and hooks in anterior<br />
setigers are <strong>of</strong>ten lighter in color <strong>the</strong>n those far<strong>the</strong>r back, and<br />
smaller specimens tend to have more light-colored aciculae<br />
than do larger specimens.<br />
Cladistic Analysis<br />
Species <strong>of</strong> <strong>Eunice</strong> have for <strong>the</strong> last 50 years organized into<br />
informal species groups (Hartman, 1944; Fauchald, 1970). A<br />
cladistic analysis was performed to test if <strong>the</strong>se informal groups<br />
represented monophyletic, possibly definable taxa. The analysis<br />
was done using PAUP 2.4 on an IBM PS-2/80 and <strong>the</strong><br />
corresponding mainframe version on an IBM 4381.<br />
SELECTION OF OUTGROUP TAXA<br />
This initial analysis was intended to justify selection <strong>of</strong><br />
outgroups to be used in this study. A complete analysis <strong>of</strong> <strong>the</strong><br />
order Eunicoidea is beyond <strong>the</strong> scope <strong>of</strong> this study. The