On the junior subjective synonyms of Coullia Hamond , 1973 ( Copepoda , Harpacticoida , Laophontidae ) : an update and key to species and related genera

Morphological comparison of the genera Coullia Hamond, 1973 and Phycolaophonte Pallares, 1975 (Copepoda, Harpacticoida, Laophontidae) shows that the latter should be relegated to a junior synonym of the former. Th e forgotten generic name Eolaophonte Apostolov, 1990, introduced for two species displaying the plesiomorphic condition of P2–P4 endopodal segmentation, is also considered synonymous with Coullia since the taxon it denotes lacks a distinct apomorphy, rendering it paraphyletic exclusive of Coullia. Pesta’s (1959) record of Laophonte sp. from Sorrento (Italy) pertains to Coullia mediterranea (Apostolov, 1990) comb. nov. Keys to the six species currently included in Coullia and to the laophontid genera displaying endopodal size discrepancy (P2 endopod smallest) are presented.


Introduction
A recent review of the outstanding nomenclatural problems in the Harpacticoida (Huys in press) resulted in the discovery of a new genus-group name proposed in 1990, which had escaped the attention of recent compilers (Bodin 1997, Boxshall and Halsey 2004, Wells 2007) and is not indexed in the Zoological Record.Apostolov (1990) established the genus Eolaophonte (family Laophontidae) for a new species, E. mediterranea Apostolov, 1990, based on two adult females recovered from algal washings in Monaco, and for Laophonte ?platychelipusoides Noodt, 1958, which was designated as the type species.Mor-phological comparison provides compelling evidence that both Eolaophonte and Phycolaophonte Pallares, 1975 are junior subjective synonyms of Coullia Hamond, 1973.Noodt (1958) described Laophonte ?platychelipusoides based on a single female from Tenerife.He suspected a relationship with Lang's (1948) inopinata-group in the genus Laophonte Philippi, 1840 but the lack of information on the male made him consider this assignment provisional and rank it instead as species incertae sedis in the Laophontidae.Noodt also claimed similarities with Hemilaophonte Jakubisiak, 1932 and remarked on the convergent swimming leg morphology in the genus Platychelipus Brady, 1880 (hence his choice of the specifi c name).Conversely, Lang (1965) believed L. ?platychelipusoides could be assigned to either Paralaophonte Lang, 1948 or Arenolaophonte Lang, 1965 but this suggestion was based solely on (unspecifi ed) female characters.Hamond (1973) proposed the genus Coullia for a new deepwater species from the North Carolina continental shelf, C. heteropus Hamond, 1973 (also known from one female only), and assigned L. ?platychelipusoides to this genus.He considered Coullia unique in its morphology of P2-P4, having no inner setae on the exopods and, more signifi cantly, reduced endopods with that of P2 being smaller than the others (in virtually all other Laophontidae with endopodal size discrepancy known at the time it was the P4 that was smallest).His claim that the absence of terminal setae on P2-P4 exp-3 is also a generic diagnostic is unfortunately based on an oversight; the inner apical setae are present but minute and often obscured by the large outer distal spines.It should be noted that the spelling platychelipusioides, fi rst introduced by Lang (1965) and subsequently adopted by other authors (e.g.Hamond 1973, Fiers 1992a, Bodin 1997, Gómez and Boyko 2006), is to be considered an incorrect subsequent spelling (ICZN Art. 33.3).Apostolov (1990) used the endopodal segmentation of P2-P4 to restrict the generic concept of Coullia (to species exhibiting 1-segmented endopods) and to justify the proposal of his new genus Eolaophonte (species with 2-segmented endopods).To the latter he removed C. platychelipusoides, which he designated as the type species, and a new species E. mediterranea, from Monaco.In the former he grouped Coullia heteropus and Hemilaophonte clysmae Por & Marcus, 1973, originally described by Por and Marcus (1973) from the Suez Canal.Fiers (1992a), who, like other authors, was not aware of Apostolov's (1990) paper, also removed H. clysmae to the genus Coullia.Comparison of Apostolov's (1990) diagnoses shows that the only notable diff erence between Eolaophonte and Coullia is the plesiomorphic expression of the small proximal endopod segment in P2-P4 in the former, thus rendering Eolaophonte a paraphyletic taxon.Pending the discovery of apomorphic character states which may characterize the taxon, Eolaophonte must be relegated to a junior subjective synonym of Coullia.Pesta (1959) illustrated the P5 and caudal rami of a damaged female from a submarine cave near Sorrento (Italy), which he tentatively identifi ed as Laophonte sp.Although he alluded to similarities with Laophonte inopinata T. Scott, 1892 in the caudal ramus, and to Heterolaophonte curvata (Douwe, 1929) in the P5, it is now more than conceivable that he was dealing with Eolaophonte (= Coullia) mediterranea.Gómez and Boyko (2006) admitted diffi culties in maintaining Coullia and Phycolaophonte as distinct genera, and the combination of characters (i.e.shape and armature of caudal ramus and female P5, 6-segmented antennule, armature and shape of P2-P4) used by these authors to place their new species Phycolaophonte tongariki Gómez & Boyko, 2006 is not exclusive to the latter genus.Consequently, Phycolaophonte is here subsumed into the synonymy of Coullia since there is no fundamental diff erence between its type species P. insularis (and P. tongariki) and species previously assigned to Eolaophonte.Pallares (1975a) herself had already hinted at a relationship with Coullia heteropus.Th e genera Phycolaophonte, Eoalophonte and Coullia collectively represent a monophyletic lineage in which each of the three "genera" merely illustrate diff erent stages in the gradual reduction in swimming leg setation and segmentation (Table 1).

Results and discussion
Other, as yet undescribed, species of Coullia have been reported in washings of Maja squinado (Herbst, 1788) from the Mediterranean and of unidentifi ed decapods from the Eastern Pacifi c (Fiers 1991(Fiers , 1992a)), in washings of the algae Hypnea musciformis (Wulfen) and Gracilaria foliifera (Forsskål) in South Carolina (Coull et al. 1983) and in the 'aufwuchs' communities of marine macrophytes in New Zealand (Hicks 1977, Coull andWells 1983).

Key to the species of Coullia Hamond, 1973
Th e key to species below should be used with caution since several, as yet undescribed, species are known to exist (Hicks 1977, Coull and Wells 1983, Coull et al. 1983, Fiers 1991, 1992a).Secondly, there is considerable confusion in the literature with regard to the exact setal formulae for the P2-P4 (particularly the endopods) as several minute elements have almost certainly been overlooked or tube-pores may have been misinterpreted as rudimentary setae.Finally, little is known about the sexual dimorphism expressed in the swimming legs since only the male of C. insularis has been formally described (Table 1 Additional notes.In addition to the type locality, Por and Marcus (1973) found C. clysmae also in algal washings at Port Taufi q, outside the Suez Canal (Gulf of Suez).
Table 1.Armature formulae of Coullia species.Th e setal formulae of P2-P4 exp-3 have been corrected for the minute inner apical setae which may have been overlooked in some species [they are consistently present in detailed species descriptions (e.g.Mielke 1985, Gómez andBoyko 2006)]; it is possible that they are genuinely absent on P4 exp-3.a Por and Marcus (1973) claimed P3-P4 endopods have 3 setae; their fi gure of the P3 (Fig. 39) is inconclusive in this respect and requires confi rmation; the tiny element between the 2 setae in both P3 and P4 may represent the apical tube-pore found in most other species (see e.g.Mielke (1985) for P4).b According to Pallares (1975a) some specimens have only 2 outer spines (as in the ♂).Fiers (1992a) noted that the text and fi gures describing the P2 and P3 are contradictory; Fig. 40 in reality illustrates the P2, not the P3, and vice versa.Coullia clysmae diff ers from its congeners in the complete absence of the P2 endopod; this character, and the loss of the outer spine on P1 exp-1 require confi rmation.Pallares (1975a) described C. insularis from washings of Macrocystis pyrifera (L.) and Delesseriaceae (red algae) collected in Bahía Vancouver, Isla de los Estados (Argentina).In a subsequent paper, Pallares (1975b) recorded the species from the plankton surrounding Macrocystis beds and washings of various algae including Durvillea and Delesseriacea.Mielke (1985) added a second record from Maiquillahue, central Chile, and updated the description of the male.Th e only notable diff erence between the two populations is found in the caudal ramus which appears more slender, having concave margins and a shorter seta II (as long as seta I) in the Argentinian material.Mielke remarked that Pallares (1975a) had reversed the outer and inner margin of the caudal ramus in her text description.Pallares noted variability in the number of outer spines on P4 exp-3 in the ♀ (2 or 3) but not in the ♂ (2); if 3 spines is the normal condition it implies that the P4 exopod is sexually dimorphic in those species that have retained that number in the female.Th e sexual dimorphism on the exopods of P2-P3 (exp-2 with inner seta) is probably diagnostic for all species of the genus since it is also expressed to a certain extent in the related genera Robustunguis and Psammoplatypus (Lee and Huys 1999).
Coullia mediterranea shares with C. clysmae the minute and unarmed proximal exopod segment of P1.It is unclear whether this indicates common ancestry or is merely the result of imperfect observation.Pesta's (1959) record of this species is based on a single female abdomen found in a submarine cave in the Gulf of Sorrento (Italy).Pesta gave illustrations of the P5 and caudal rami.Based on the apically recurved caudal seta V he claimed a certain similarity with L. inopinata, a species not yet recorded from the Mediterranean, but also admitted that the diff erence in leg 5 setation probably indicated that the specimen collected was juvenile.
Th e remaining three species (platychelipusoides, heteropus, tongariki) are known from their respective type localities (Tenerife, North Carolina, Easter Island) only.Noodt (1958) described the female antennule of C. platychelipusoides as indistinctly 8-segmented with a partial suture subdividing the apical segment, however, Lee and Huys (1999) reinterpreted or re-examined the 8-segmented condition reported for some species of Paralaophonte and Heteronychocamptus Lee & Huys, 1999 and concluded that the ancestral state for the family Laophontidae is 7-segmented.Coullia heteropus is thus far unique within the genus by the presence of only three elements on the female P5 baseoendopod.Hamond (1973) overlooked the minute inner distal seta on P2-P3 (and possibly P4) exp-3, and the pinnate ornamentation on the exopodal spines of P4.