GONZALES & GOSLINER: NEW SPECIES OF PHILINE FROM THE INDO-PACIFIC 379 
in R multipapillata. Philine multipapillata has a distinct genital ganglion present on the visceral 
loop of the central nervous system, whereas it appears to be absent in P. habei, P puka, and P. ver- 
densis. It has not been described in P mbrci. The penial papilla of P verdensis is hammer-shaped 
with a prominent flap on the anterior lobe, whereas in P habei the penial papiUa is asymmetrical 
with one lobe larger than the other or more of less equally lobed (present study). The penis of 
P piikah smaller with markedly unequal lobes and a veiy short base. None of these have a large 
penial papilla with dense conical papillae found in P multipapillata. The shape of the penial papil¬ 
la remains unknown for P rubra. Based on these differences, it is clear that P. multipapillata is dis¬ 
tinct from all Other species described within the P. aperta clade. 
Phylogenetic Analysis 
Our molecular phylogenetic analysis includes five species studied here plus P. habei, P. area, 
and P. rubrata and those studied by Krug et al. (2012) and six other taxa from GenBank (Kmg et 
al. 2012).The resulting maximum likelihood tree (using RAxML) molecular phylogeny is depict¬ 
ed in Figure 14. 
In our analysis, Philine verdensis is most closely related to P. habei, but that sister group rela¬ 
tionship is not strongly supported. The two species have a p-distance of just over 4% (Table 2) for 
the 16S gene, strongly supporting their distinctness as separate species. Currently, there are no 
molecular data for P puka and P. rubra, the other two species with large pores on the gizzard plates. 
Together, P. verdensis and P. habei form a weakly supported sister-group relationship with 
P. quadripartita, P. aperta, P angasi, P paucipapillata, and an undeseribed species from Australia. 
Philine dentiphallus appears as sister species to P. auriformis and P pittmani is sister to both of 
these species, although these relationships are relatively weakly supported. Philine acuticauda is 
sister to P. hearstorum in our analysis and this relationship is strongly supported. Together these 
species forni the sister group to the Philine aperta clade, but that relationship is weakly supported. 
The sister group to these Philine species includes seven species of aglajids and Philine orca and 
P. rubrata. The relationships within this clade are weakly supported. 
Table 2. Minimum 16S pairwise uncorrected p-distance among Philine species described in this paper. 
habei 
verdensis 
pittmani dentiphallus 
acuticuada 
hearstorum 
habei 
verdensis 
0.042 
pittmani 
0.121 
0.116 
dentiphallus 
0.160 
0.122 
0.085 
acuticauda 
0.169 
0.147 
0.137 0.166 
hearstorum 
0.168 
0.152 
0.145 0.178 
0.027 
auriformis 
0.105 
0.104 
0.072 0.067 
0.155 
0.162 
Discussion 
This work includes the description of six new species of Philine from the tropical Indo-Pacif- 
ic. Both morphological and molecular data support the separation of these species. Four of the six 
species are members of the Philine aperta clade, a group that is already been shown to be well rep¬ 
resented in the Indo-Pacific tropics (Price et al. 2011). The other two species, P. acuticauda and 
P. hearstorum, have the same radular morphology and gizzard plate morphology as P. alba Mat¬ 
tox, 1958, from the Pacific coast of California and P. alboides, from the Caribbean. The two new 
species represent the first members of this group known from the tropical Indo-Pacific. 
Krug et al. (2012) conducted the first molecular phylogeny of Philine, but only included Phi- 
