352 
THE CORAL TRIANGLE: HEARST BIODIVERSITY EXPEDITION 
Many morphological descriptions of Philine species have been undertaken over the past sev¬ 
eral decades (Marcus 1974; Marcus and Marcus 1966, 1967, 1969; Rudman 1970, 1972a, b; 
Gosliner 1988), and these studies describe new taxa and review the anatomy of previously identi¬ 
fied species. Price et al. (2011) also provided the first morphologically-based phylogenetic study 
of Philine. More recently, Krug et al. (2012) provided a molecular phylogeny based on the 16S 
mitochondrial gene, that was used primarily to establish the origin of two introduced species of 
Philine now found on the Pacific coast of North America. Ohnheiser and Malaquias (2013) 
reviewed the North Atlantic Philine species, described two additional species from the region and 
presented a molecular phylogeny of these species based on the COI mitochondrial gene. 
The World Register of Marine Species (WORMS: <http\//www.mannespecies.org/ 
aphia.php?p^taxdetails&id= 138339>) lists 117 valid species of Philine from throughout the 
world’s oceans. As noted by Price et al. (2011), the majority of these are known only from shells 
and their identity remains obscure. The Indo-Pacific tropics and adjacent temperate regions support 
a diversity of philinid opisthobranchs (Gosliner et al., 2008), with 24 described species document¬ 
ed from the region (Price et al. 2011, WORMS). Several relatively recently described species have 
been added to the Indo-Pacific region since the systematic reviews cited above (Valdes 2008; Price 
et al. 2011). These include Philine abyssicola Valdes, 2008, P babai Valdes, 2008, P habei Valdes, 
2008, P fenestra Price, Gosliner and Valdes, 2011, R paucipapillata Price, Gosliner and Valdes, 
2011, P puka Price, Gosliner, and Valdes, 2011 and P. sarcophaga Price, Gosliner and Valdes, 
2011. Krug et al. (2012), illustrated that at least one additional Indo-Paeific species from Australia 
as unidentified and quite likely undescribed. 
Recent biodiversity exploration of deeper waters surrounding the Philippines from the 2004 
Panglao hiternational Expedition and the 2011 Hearst Philippine Biodiversity Expedition has pro- 
dueed additional material, wliich is the focus of the present study. Examination of this material plus 
specimens of a shallow-water species from the Hawaiian Islands reveals the presence of six addi¬ 
tional undescribed species. The description of these species and the examination of their molecu¬ 
lar phylogenetic relationsliips formulate the focus of the present review of the genus Philine. In 
order to compare these species with other species known from the Pacific and Indo-Pacific, our 
molecular study employs the examination of the 16S gene studied by Krug et al. (2012) rather than 
using the COI gene used by Ohnheiser and Malaquias (2013) for Atlantic taxa. The intent of the 
present molecular study is to detennine the distinctness of the species described here, as well as 
further assessing the phylogeny and monophyly of Philine. 
Materials and Methods 
Morphological Methods 
Over the last decade, new specimens of philinds were collected from the Verde Island Passage 
and Bohol Island, Philippines and the Hawaiian Islands. Specimens collected were preserved in 
either 10% formalin for proper preseiwation of anatomical stmctures, or preserved entirely in 95% 
ethanol for later moleculai* study. Prior to preservation of any specimen in formalin, a tissue sam¬ 
ple was taken from the animal and preserved in 95% ethanol for later moleeular study. 
Upon return to the California Aeademy of Sciences, dissections were completed, and drawings 
of anatomical structures were accomplished using a Nikon SMZ-U binocular microscope with 
drawing tube. Buccal mass structures of specimens were dissected and cleaned by placing them in 
a 10% NaOH solution for 4—10 hours, and then prepared for scanning elecfron microscopy. The 
radula was separated and cleaned of any remaining tissue. It was then placed on a glass cover slip, 
air-dried, and then mounted on a stainless steel stub. The same procedure was followed for exam- 
