120 



J.D. TAYLOR AND E.A. GLOVER 



Table 2 Matrix of Cardiolucina species and the outgroup Parvilucina 

 tenuisculpta and their character states. Character coding as in Table 1 . 





1 



2 



3 



4 



5 



6 



7 



8 



9 



10 



11 



12 



agassizi 







1 







1 







1 



1 



















1 



australopilula 









2 







1 







1 



1 











2 



1 



civica 









2 



1 







1 



1 



1 











1 



2 



crassilirata 









2 



1 







1 



1 



1 











1 



2 



eucosmia 





1 



1 







1 



















1 











pisiformis 

















1 















1 















quadrata 









2 



1 







1 



1 



1 











2 



1 



rugosa 







1 







1 







1 















1 







1 



semperiana 







1 







1 







1 















1 







1 



siquijorensis 



1 















1 















1 



? 











lamothei 







2 







1 







1 























2 



rehderi 







2 







1 







1 























1 



P. tenuisculpta 



2 







3 



1 







2 



1 



















2 



original designation by Dall (1901), Cardiolucina (as Bellucina) has 

 been widely listed as a subgenus of Parvilucina and the shells have 

 some features in common (descriptions in Hickman 1994). More- 

 over, there are some similar anatomical features, P. tenuisculpta 

 having a visceral lobe resembling that found in C. semperiana and 

 C. australopilula, as well as the connection of the posterior ctenidia 

 to the mantle and the lack of a fused inhalant aperture (Reid and 

 Brand, 1986). The characters and their states are listed in Table 1. 



The distribution of the various character states amongst the 

 Cardiolucina species and the outgroup is shown in Table 2. 

 Phylogenetic analysis using this matrix produced 2 equally parsimo- 

 nious trees of 26 steps with a consistency index of 0.731 and a 

 homoplasy index of 0.269. A strict consensus tree is shown in Fig. 

 29. The differences in the 2 trees resulted from the grouping of 

 C. civica and C. crassilirata as a single clade or as a trichotomy with 

 the C. australopilula IC. quadrata clade. 



The analysis suggests that there are three distinct clades of 

 Cardiolucina species (Fig. 29). Clade 1 contains C australopilula. 



crassilirata, civica and quadrata. Apomorphies which define this 

 clade at Branch 2 are; the tiny lunule, the hooked- shaped, anterior 

 cardinal tooth and the hinge buttress. Clade 2, separated at Branch 4 

 by a single apomorphy of having less than 20 concentric lamellae, 

 contains two major branches; that comprising the type species C. 

 agassizi and the species around C. semperiana. (Branch 7) and 

 another branch (Branch 5) containing C. eucosmia, pisiformis and 

 siquijorensis. This latter clade is defined by four apomorphies; 

 having less than 20 radial ribs, which are straight-sided and cross the 

 concentric lamellae and also possess coarse marginal crenulations. 

 The C. agassizi - lamothei clade is defined at Branch 7 by the 

 prominent posterior sulcus and a broad anterior sulcus. 



This hypothesis of relationships should be regarded as tentative 

 because it is based on shell characters alone. Nevertheless, it is open 

 to falsification by additional or alternative character sets. 



GEOGRAPHICAL DISTRIBUTION 



Two species, Cardiolucina lamothei and C. rehderi are found on 

 opposite sides of the Atlantic, whilst the other nine are distributed 

 largely within the tropical Indo-West Pacific Province. One of these, 

 C. semperiana, has a wide range from the Gulf of Suez to Philip- 

 pines whilst the rest have more restricted distributions within the 

 central Indo-Pacific. Four species are endemic to Australia; C. 

 eucosmia, C pisiformis and C. australopilula to the western and 

 northern coasts and C. crassilirata to the temperate southern coast. 

 Cardiolucina civica occurs from Japan through Indonesia to the 

 Andaman Islands in the west and Fiji in the east. C quadrata is 

 restricted to deeper water around the Philippines and Indonesia. 

 Apart from C. rehderi, which is known from only one vague locality, 

 the species with the most limited range is C. siquijorenis from 

 Siquijor Island in the Philippines. All the species are largely tropical 

 in distribution, except for C crassilirata from southern Australia. 



Table 3 Summary of shell characters in Cardiolucina species 

























Lunule 





















Sulcus 



Marginal 



small, under 



heart- 



Species 



H 



L 



T 



Radial ribs 



Concentric lamellae 



post. 



ant. 



crenulae 



beak 



shaped 



agassizi 



7.4 



7.1 



2.7 



c.25 



faint, closely 

 spaced 



c.20 



width of interspaces 

 extremely variable 



deep 



narrow 



28 





shallow 



australopilula 



13.8 



12.7 



7.5 



c.20 



interspaces almost 

 as wide as ribs 



c.34-36 



low, narrow 



shallow 



none 



21 



* 





civica 



10.6 



10.4 



4.5 



>30 



fine, closely 

 spaced 



>25 



conspicuous 



shallow 



none 



>60 



* 





crassilirata 



11.0 



11.3 



6 



>40 



fine, closely 

 spaced 



>30 



fine, clustered 

 in groups of 4-6 



shallow 



none 



31 



* 





eucosmia 



7.8 



7.7 



3.8 



c.12 



broad, with 

 wide interspaces 



c.13-14 



low 



v.deep 



narrow 



12 





v. deep 



pisiformis 



4.4 



4.4 



2 



13-17 



low, widely 

 spaced 



c.20 



narrow 



shallow 



none 



13 





deep 



quadrata 



12.9 



13.6 



5.3 



c.25 



fine,closely 

 spaced 



25-30 



low, broad 



shallow 



none 



25 



* 





rugosa 



3.1 



3.2 



1.2 



>25 



faint, closely 

 spaced 



c.20 



prominent 



deep 



narrow 



20 





shallow 



semperiana 



7.7 



7.5 



2.7 



20-30 



low, closely 

 spaced 



15-24 



interspaces variable 

 in width 



deep 

 ribbed 



narrow, 



20-21 





shallow 



siquijorensis 



3.5 



3.5 



1.3 



12 



broad 



14-16 



thin 



shallow 



none 



12 





shallow 



lamothei 



4 



3.9 



2 



>40 



fine, closely 

 spaced 



15-18 



prominent 



deep 



broad 



40 



* 





rehderi 



4.2 



4.2 



1.9 



>30 



fine, faint 

 closely spaced 



14-16 



prominent 



deep 



broad 



20 



* 





H=height, max mm, L=length, max mm, 



T=tumidity, max mm. 

















