A New Family of Harpacticoid Copepods and an Analysis of the Phylogenetic Relationships Within the Laophontoidea T

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A New Family of Harpacticoid Copepods and an Analysis of the Phylogenetic Relationships Within the Laophontoidea T Bijdragen tot de Dierkunde, 60 (2) 79-120 (1990) SPB Academie Publishing bv, The Hague Amsterdam Expeditions to the West Indian Islands, Report 64* A new family of harpacticoid copepods and an analysis of the phylogenetic relationships within the Laophontoidea T. Scott Rony Huys Marine Biology Section, Zoology Institute, State University o f Gent, K.L. Ledeganckstraat 35, B-9000 Gent, Belgium; and Delta Institute for Hydrobiological Research, Vierstraat 28, 4401 EA Yerseke, The Netherlands Keywords: Cristacoxidae fam. nov.,Cristacoxa gen. nov., Cubanocleta, Noodtorthopsyllus, Laophon­ toidea, phylogeny, Copepoda Summary (7) sexual dimorphism of P3 endopod; (8) P6 bisetose with one member fused to somite. There is no close relationship neither with the Normanellidae Lang, nor with the Ancorabolidae T. A new family Cristacoxidae is proposed to accommodate the Scott. The Laophontidae are considered the first offshoot in the monotypic genera Cristacoxa gen. nov., Noodtorthopsyllus evolution of the Laophontoidea because of the retention of the Lang (ex Canthocamptidae) and Cubanocleta Petkovski (ex 8-segmented antennule in both sexes and the ancestral seta for­ Laophontidae). Cristacoxa petkovskii gen. et spec. nov. is mulae on P2-P4. The other families can be assigned to two described on the basis of a single male collected from coralline clades: the Adenopleurellidae and the Laophontopsidae-Cris- sand of Bonaire, West Indies. N. psammophilus N oodt and C. tacoxidae-Orthopsyllidae-grouping. The Laophontopsidae and n oodti Petkovski are redescribed and refigured on the basis of the Cristacoxidae are sister groups because of the shared sexual new material from the Galápagos (Isla Santa Cruz), the West In­ dimorphism of the P3 endopod (advanced type), and the fusion dian Islands (Curaçao, Klein Curaçao, Bonaire) and the Canary of antennular segments distal to the geniculation in the male. Islands (El Hierro, Tenerife). The new family is characterised by Emphasis is placed on the postembryonic development in the the presence of an outer spinous process on the first antennular Laophontoidea of the male P3 endopod and P6 and on their ho­ segment, the absence of the antennary exopod, the uniramous mologies in the female. A reconstruction of the hypothetical an­ mandibular palp, the presence of conspicuous cristae on the cestor of the Laophontoidea is presented. precoxa and coxa of leg 1 and the elongation of the apical exopo- dal spines of P2 to P4. The Cristacoxidae are unique in having an equal number of setae/spines on the P5 in both sexes. Exami­ nation of the ontogeny of this leg in males and females of Or­ Zusammenfassung thopsyllus spec, gives strong evidence that the cristacoxid P5 has undergone neotenous evolution. Males of Cristacoxidae are Um die monotypischen Gattungen Cristacoxa gen. nov., N o o d t­ readily recognisable by their extremely long spermatophores. orthopsyllus Lang (ex Canthocamptidae) und Cubanocleta Pet­ The new family is allocated to the superfamily Laophontoidea kovski (ex Laophontidae) unterzubringen, wird eine neue T. Scott, together with the Laophontidae T. Scott, the Laophon­ Familie Cristacoxidae vorgeschlagen. Cristacoxa petkovskii topsidae Huys & Willems, the Orthopsyllidae Huys and the gen. et spec. nov. wird auf der Basis eines einzelnen Männchens Adenopleurellidae Huys. A phylogenetic analysis of the rela­ beschrieben, das von Korallensand auf Bonaire, Westindische tionships within the superfamily is presented, and as a result Inseln, gesammelt wurde. N. psammophilus Noodt und C. Por’s (1986) concept of the Laophontoidea is refuted. Instead, n oodti Petkovski werden anhand neuen Materials von Galápa­ it is concluded that the superfamily can be defined on the basis gos (Isla Santa Cruz), den Westindischen Inseln (Curaçao, Klein of the following apomorphies: (1) antennules with outer spinous Curaçao, Bonaire) und den Kanarischen Inseln (El Hierro, process on segment 2; (2) antenna with allobasis bearing 1 seta; Teneriffa) neu beschrieben und neu illustriert. Die neue Familie (3) antennary exopod quadrisetose; (4) PI exopod without inner ist charakterisiert durch das Vorhandensein eines äußeren seta on exp-2 and 4 setae/spines on exp-3; (5) PI endopod Dornfortsatzes am ersten Antennulasegment, das Fehlen eines 2-segmented with elongated enp-1 and 2 elements on enp-2; (6) Exopoditen an der Antenna, den uniramen Mandibularpalpus, P2-P4 with 2-segmented endopods; exp-1 without inner seta; das Vorhandensein auffälliger Leisten (Cristae) an der Praecoxa * Report 63 will be published in Beaufortia vol. 41 (1990). Contribution No. 455 of the Delta Institute for Hydrobiological Research, Yerseke. i 80 R. Huys - A new family of harpacticoid copepods und Coxa von P 1 und die Verlängerung der apikalen Dornen an (1955). This remarkable copepod, described from a den Exopoditen von P2-P4. Die Cristacoxidae sind einmalig coarse sandy beach on Tenerife, differs profoundly dadurch, daß P5 in beiden Geschlechtern die gleiche Anzahl von from the other species in many aspects (e.g. spinous Borsten/Dornen trägt. Die Überprüfung der Ontogenie dieses Beines bei männlichen und weiblichen Individuen von Ortho­ process on different segment of antennule, absence psyllus spec, macht es sehr wahrscheinlich, daß P5 in den of antennary exopod, ornamentation of swimming Cristacoxidae eine neotene Entwicklung durchgemacht hat. legs, P5 in both sexes) but Noodt (1955) attributed Männliche Cristacoxidae sind leicht an ihren extrem langen these differences to the “... stark abgeleitete und Spermatophoren zu erkennen. Die neue Familie ist zusammen daher isolierte Stellung ...” of the species. Lang mit den Laophontidae T. Scott, Laophontopsidae Huys & Wil­ lems, Orthopsyllidae Huys und den Adenopleurellidae Huys der (1965) rightly concluded that O. psammophilus Superfamilie Laophontoidea T. Scott zugeordnet. Eine phylo­ must have been splitted off before the evolution of genetische Analyse der Beziehungen innerhalb der Superfamilie the other Orthopsyllus- species, because the struc­ wird vorgenommen und als deren Ergebnis Por’s (1986) Konzept ture of the antennule and the armature of the sec­ der Laophontoidea widerlegt. Statt dessen wird geschlußfolgert, ond to fourth legs cannot be derived from the con­ daß die Superfamilie anhand folgender Apomorphien definiert werden kann: (1) Antennula mit äußerem Dornfortsatz am 2. ditions found in the other members of the genus. Segment; (2) Antenne mit Allobasis, die eine Borste trägt; (3) Lang (1965) therefore proposed a new genus, Exopodit der Antenne mit vier Borsten; (4) Exopodit von PI Noodtorthopsyllus, to accomodateO. psammophi­ ohne innere Borste an exp-2 und vier Borsten/Dornen an exp-3; lus as its type and only species. The new genus and (5) Endopodit von PI zweigliedrig mit verlängertem enp-1 und its presumed relative Orthopsyllus were kept in the zwei Anhänge an enp-2; (6) P2—P4 mit zweigliedrigen Endo- poditen; exp-1 ohne innere Borste; (7) Sexualdimorphismus bei Canthocamptidae until Huys (in press c) removed Endopit P3; (8) P6 mit zwei Borsten, ein Bein mit dem Körper­ the latter genus to a separate family. segment verschmolzen. Weder zu den Normanellidae Lang noch The closest relative of N. psammophilus is zu den Ancorabolidae T. Scott gibt es enge Beziehungen. Die without doubt Cubanocleta noodti Petkovski Laophontidae werden wegen der Beibehaltung der achtgliedri- which is described from the mouth of the Rio B aco­ gen Antennula in beiden Geschlechtern und der ursprünglichen Borstenformel von P2-P4 als erste Abzweigung in der Evolu­ nae in Cuba (Petkovski, 1977). According to Pet­ tion der Laophontoidea angesehen. Die anderen Familien kön­ kovski the absence of an antennary exopod com­ nen zwei Linien zu geordnet werden: den Adenopleurellidae und bined with the structure of the rostrum, the der Laophontopsidae-Cristacoxidae-Orthopsyllidae-Gruppe. thoracopods and the caudal rami suggests that C. Die Laophontopsidae und Cristacoxidae sind Schwestergruppen noodti represents a link between the Ancorabolidae wegen des bei beiden vorhandenen Sexualdimorphismus des Endopodits von P3 (fortgeschrittener Typus) und der an der An­ and the laophontid subfamily Normanellinae. He tennula des Männchens distal von der Biegung vorhandenen further recognised also a certain resemblance with Verschmelzung der Segmente. Besondere Bedeutung wird bei some odd genera (Laophontella Thompson & A. den Laophontoidea der postembryonalen Entwicklung des En­ Scott; Apolethon Wells) of which the affinity to the dopodits von P3 und von P6 des Männchens und der homologen Laophontidae was or is still doubtful. Teile beim Weibchen beigemessen. Eine Rekonstruktion des hypothetischen Vorfahren der Laophontoidea wird vorgestellt. A third closely related species was collected by Prof. Dr. J.H. Stock in coralline sand of Bonaire, Introduction Netherlands Antilles and is described herein as a new genus Cristacoxa gen. nov.. N. psammophilus The history of the genusOrthopsyllus Brady & and C. noodti are redescribed on the basis of Robertson marks the course of a gradual recogni­ material collected in the Galápagos (Isla Santa tion of the high variability in many of its morpho­ Cruz), the West Indian islands (Curaçao, Klein logical characteristics (Lang, 1965; Wells, 1968; Curaçao, Bonaire) and the Canary Islands Hamond, 1970) and the process of lumping to­ (Tenerife, El Hierro). Despite their laophontoidean gether geographically widely separated populations relationships it is concluded that the three genera into
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