BASAL LAOPHONTID EVOLUTION 



103 



Table 4. States for characters listed in Table 3 [0 = plesiomorphic; 1 = apomorphic; 2 = further derived state]. Characters 14 and 22 are multistep 

 characters. 



Taxon 



2 3 4 5 6 7 



9 10 11 12 13 14 15 16 17 



19 20 21 22 23 24 25 



ARCH1LAOPHONTE 



MOUREPHONTE 



bulbifera 



bulligera 



galapagoensis 



hirsuta 



longicauda 



longiremis 



spelaea 



typhlops 



canalis sp. nov. 



compacta sp. nov. 



hamondi sp. nov. 



lobata sp. nov. 



profunda sp. nov. 



vervoorti sp. nov. 



spec, sensu Chislenko (1967) 



spec, sensu Mielke (1997) 



Laophonte cornuta-group 



Onychocamptus 



OTHER LAOPHONTIDAE 











1 

 1 





 



1 





 1 

 1 





 7 



1 







9 



1 



1 



9 



1 









1 

 1 





 

 1 









 



1 



7 

 

 1 





 1 



1 







1 

 1 



1 

 1 

 1 





 

 







1 

 1 







9 









 

 





 

 





 7 



1 









 

 



























1 





1 

















1 



1 



1 







1 















1 











9 



7 

 1 

 1 



7 

 1 

 1 



1 

 1 

 1 



9 

 



7 



1 







1 



1 

 1 

 1 





7 

 1 

 1 



1 





 1 





1 







1 



7 

 



1 



1 

 1 

 ? 



1 



1 

 1 



1 

 1 

 1 



1 







1 



1 

 1 

 1 







1 









 

 





 

 7 



1 

 1 

 1 







 

 





















7 













7 

















7 



1 











? 







9 



7 









1 





7 



7 











7 











7 



9 



7 





7 



7 



7 



7 











7 











1 



1 















1 







1 







1 





 1 







1 







1 







1 



9 



1 

 1 





 1 







 7 





 1 



1 

 1 



1 

 1 







1 





 







1 





 



1 



7 







9 







1 





1 









1 





























1 









9 







1 



1 











1 







9 



9 











































9 



1 

















1 



1 



















7 



? 









1 





7 



7 



7 

 1 

 1 



1 

 1 

 1 



7 

 1 

 



1 

 1 



1 



1 

 1 

 1 



9 



1 

 1 



1 



1 

 1 

 1 





7 

 7 

 1 



7 



7 

 1 

 1 



7 

 1 



1 



1 

 1 

 1 





 

 



1 

 1 

 1 





 

 







7 









 7 







1 

 1 

 1 







 

 





7 



? 



7 







? 



7 



7 





7 













7 



7 



? 











7 







1 







7 





1 



7 



1 



7 



7 



1 



7 



1 



1 



7 



1 



1 





7 





7 



7 



1 



1 



1 







7 



7 



1 









1 



























1 

























1 



2 



1 







1 



1 



















































1 















1 















2 



1 







1 



1 











2 





































































1 











1 



1 







1 



















further elaborated in both Mourephonte and the residual species of 

 Esola by the development of an accessory pair of anteroventral 

 pores on the cephalothorax (character 2) and of ventral or medial 

 pores on the caudal rami (character 4). Finally, the lateral pores on 

 the genital (double-)somite (character 3) evolved not until after the 

 divergence of Mourephonte. 



The genus Esola is redefined here to encompass the terminal 

 polychotomy containing the type species E. longicauda and 7 other 

 species (Fig. 32). This strongly supported, cosmopolitan crown- 

 group is characterized by distinctive labral ornamentation, caudal 

 ramus sexual dimorphism, formation of 3 spinous processes on the 

 first antennulary segment and modification of segment 5 in the male 

 antennule E. hirsuta is the only species that shares genital cup- 

 shapes pores with this clade, however it is excluded from Esola and 

 placed in a monotypic genus Applanola on account of the following 

 autapomorphies: (1) dorsoventrally depressed body morphology, 

 (2) elongation of mandibular palp, (3) modification of PI endopod, 

 (4) exopodal sexual dimorphism of P2-P3, (5) loss of outer spine on 

 P2 enp-2, and (6) strong reduction of the male sixth legs. The sexual 

 dimorphism on the P2-P3 exopod is unique in the Esolinae. Al- 

 though this character is globally homoplastic within the Laophontidae 

 it can be informative locally (see Lee & Huys, 1999) and should not 

 therefore be routinely ignored in phylogenetic analyses. 



The three remaining species, compacta, spelaea and bulligera, 

 are identified as independent lineages splitting off successively 

 between the basal Archesola clade and the terminal ({Mourephonte 

 + Esola) + Applanola) clade. Corbulaseta gen. nov., accommodat- 

 ing E. bulligera, is most closely related to the latter clade because of 

 shared fusions in the female antennule (segments 6—7) and PI 

 exopod (exp-2 and -3). The modified distal inner seta forming a 

 trapping-basket is a unique autapomorphy for this genus. The 

 position of Troglophonte is tentative pending the confirmation of 

 cup-shaped pores on the cephalothorax and of the armature patterns 

 of P2 exopod and P5 in both sexes. The basal position of the genus 

 Bathyesola is caused by its retention of the maximum number of 

 setae on the female P5 baseoendopod. 



The genus Mourephonte is radically divergent from other esolinids. 

 The extreme development of the PI, the complete absence of the P2 

 endopod, the loss of the inner seta on the P2-P4 exopods and the 



wide separation of the apical setae on P4 enp-2 form a remarkable 

 combination of autapomorphies which places it on a distinct evolu- 

 tionary lineage, ruling out possible inclusion in the genus Esola 

 under a broader concept. 



The residual laophontids, comprising 95% of the known species, 

 are grouped in the subfamily Laophontinae. All 54 genera have lost 

 the inner seta on PI enp- 1 and the outer spine on P2 enp-2, and bear 

 a maximum of 2 setae on the maxillipedal syncoxa (absence of 

 proximal seta). With the exception of the genus Onychocamptus and 

 the Laophonte cornuta-group all Laophontinae are characterized by 

 the P3 endopod sexual dimorphism involving the loss of the proxi- 

 mal inner seta of enp-2 (character 21). The isolated position of the 

 cornuta-gToup (= Laophonte Group I + adduensis + ciliata: Table 2) 

 testifies to the widely accepted polyphyletic status of the genus 

 Laophonte and has major nomenclatural consequences because of 

 its inclusion of the type species L, cornuta Philippi. Restriction of 

 the generic concept to the cornuta-group will require the other 37 

 species of Laophonte to be re-allocated to other existing or new 

 genera. This is a major task which can only be accomplished by 

 sound phylogenetic analysis involving the remaining laophontinid 

 genera. The sistergroup relationship between the cornuta-group and 

 Onychocamptus depicted in Fig. 32 is not be taken as absolute since 

 other advanced but closely related genera such as Folioquinpes have 

 deliberately been omitted from the outgroup to the Esolinae. Al- 

 though inclusion of these genera in future analyses may introduce 

 additional basal nodes changing the relative position of Laophonte 

 and Onychocamptus, we envisage that the latter will consistently 

 show up as an early speciation event predating the evolution of the 

 other Laophontinae. 



Subfamilial division 



ESOLINAE subfam. nov. 



Rostrum delimited at base by surface suture; antennule 9 6- or 7- 

 segmented, usually without spinous process on segment 2 but 

 frequently with processes on segment 1 ; 7-segmented and haplocerate 

 or subchirocerate in 6*, with only 2 segments distal to geniculation; 



