GONZALES & GOSLINER: NEW SPECIES OF PHILINE FROM THE INDO-PACIFIC 363 
genital atrium, where it joins the elongate duct of the bursa copulatrLx. The bursa is large and spher¬ 
ical. Its duct is narrow where it joins the bursa and widens and curves until its widest portion at the 
genital atrium. From the genital atrium the open, ciliated spenn groove leads to the cephalic penis. 
The penis (Fig. 4F, 5C) consists of a thick penial sac and an elongate, unbranched, anteriorly direct¬ 
ed prostate gland that is joined to the penial sac by a narrowed constriction. Within the penial sac 
is an elongate, lobed penial papilla that is devoid of any armature. 
Remarks. — Price et al. (2011) described a series of plesiomorphic characteristics found in 
species of Philine. Philine hearstorum has several relatively plesiomorphic features including a rel¬ 
atively large radula, more than one outer lateral tooth in the radular ribbon, a suprainestinal gan¬ 
glion situated posterior to the circumesophageal nerve ring and a simple, unarmed penial papilla 
with a single unbranched prostate. This species also has an elongate posterior end of the posterior 
shield. 
Philine hearstorum is most similar in its morphology to P, alba Mattox, 1958, from the south¬ 
ern California coast, P. alboides Price, Gosliner, and Valdes, 2011, from the Gulf of Mexico and 
Caribbean and P acuticauda, with which P hearstorum is sympatric. No molecular data are avail¬ 
able for P alba and P alboides, since all material is either fixed in formalin or the alcohol speci¬ 
mens are very old and there was no amplification of DNA (P. Krug, pei*sonal communication). The 
j9-distance (Table 2) between P, acuticauda and P. hearstorum for the 16S gene is 2.7%. This is 
consistent with the percentages found by Kmg et al. (2012) for distinct species of Philine, Moipho- 
logically, these four species have a lai*ge body size with a broad body profile and a broad, smooth, 
flattened shell with an expanded posterior wing. They also have a large radula with a wide inner 
lateral tooth that either lacks denticles or has gi'eatly reduced denticles. All four species have two 
outer lateral teeth and a simple penis and prostate. They all have a supraintestinal ganglion locat¬ 
ed posteriorly to the nerve ring. Despite these similarities, P hearstorum differs significantly from 
these taxa in several regards. It is the only species that has distinct sculpture on the shell. The pos¬ 
terior end of the body of P alba and P alboides is blunt rather than being acutely tapered as in 
P. hearstomm and P. acuticauda. The posterior end of the body is more acute in P. acuticauda. In 
P. acuticauda, P, hearstorum, and P, alboides, the inner lateral teeth entirely lack denticles, while 
in P. alba, some of the teeth have minute denticles on the masticatory border, while other teeth in 
the same radula may entirely lack denticles. In P. hearstorum, P. alba, and P. alboides, the two 
outer lateral teeth ai*e similar in shape with evenly curved cusps. In P. acuticauda, the inner of these 
two teeth is similar to those found in the other two species but the outer one has a dramatically 
reduced cusp. Philine hearstorum, P. alba, and P. alboides have three small evenly-sized gizzard 
plates, while no trace of plates was found in the two specimens of P. acuticauda examined. The 
gizzard plates of P, hearstorum have rhomboidal crystals on their surface, while those of P, alba 
and P, alboides are smooth. The prostate of P. acuticauda is only slightly curved and is posterior¬ 
ly directed, while in P. hearstomm, P alba, and P, alboides, the prostate is sharply curved, with the 
posterior end being anteriorly directed. The penial papilla of P. hearstorum has a bilobed apex, 
while the papilla of P. acuticuada, P, alba, and P. alboides is undivided. 
Philine dentiphallus Gonzales and Gosliner, sp. nov. 
Figures IE, 6-7 
Material Examined.— Holotype: body wall opened, but not dissected, CASIZ 192088, ST 
HEPD 19, beam frawl in beam trawl in muddy sand with pebbles, 541-636 m depth, western end 
of Verde Island Passage, southeast end of Golo Island, Batangas Province, Luzon, Philippines, 
13.5895°N, 120.4155°E, 1 June 2011, Hearst Expedition Deep Sea Team. Paratype: dissected. 
