Research Article |
Corresponding author: Diana M.P. Galassi ( dianamariapaola.galassi@univaq.it ) Academic editor: Saskia Brix
© 2016 Diana M.P. Galassi, Niel L. Bruce, Barbara Fiasca, Marie-José Dole-Olivier.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Galassi D, Bruce N, Fiasca B, Olivier M (2016) TA new family Lepidocharontidae with description of Lepidocharon gen. n., from the Great Barrier Reef, Australia, and redefinition of the Microparasellidae (Isopoda, Asellota). ZooKeys 594: 11-50. https://doi.org/10.3897/zookeys.594.7539
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Lepidocharontidae Galassi & Bruce, fam. n. is erected, containing Lepidocharon Galassi & Bruce, gen. n. and two genera transferred from the family Microparasellidae Karaman, 1934: Microcharon Karaman, 1934 and Janinella Albuquerque, Boulanouar & Coineau, 2014. The genus Angeliera Chappuis & Delamare Deboutteville, 1952 is placed as genus incertae sedis in this family. The Lepidocharontidae is characterised by having rectangular or trapezoidal somites in dorsal view, a single free pleonite, a tendency to reduction of the coxal plates, and the unique uropodal morphology of a large and long uropodal protopod on which the slender uropodal exopod articulates separately and anteriorly to the endopod. Lepidocharon Galassi & Bruce, gen. n. has a 6-segmented antennula, a well-developed antennal scale (rudimentary exopod), long and slender pereiopods 1–7 directed outwards, coxal plates rudimentary, incorporated to the lateral side of the sternites, not discernible in dorsal view, the single pleonite narrower than pereionite 7, scale-like elements bordering the proximal part of male pleopod 1 on posterior side, and stylet-guiding grooves of male pleopod 1 which run parallel to the outer lateral margins of the same pleopod. Lepidocharon priapus Galassi & Bruce, sp. n., type species for the genus, and Lepidocharon lizardensis Galassi & Bruce, sp. n. are described from Lizard Island, northern Great Barrier Reef. The most similar genus is Microcharon, both genera sharing the same general organization of the male pleopods 1 and 2, topology and architecture of the stylet-guiding groove of male pleopod 1, morphology of female operculum, presence of 2 robust claws of different lengths on pereiopodal dactylus 1–7, not sexually dimorphic. Lepidocharon gen. n. differs from Microcharon in the shape of the pereionites, very reduced coxal plates, the presence of imbricate scale-like elements bordering the proximal postero-lateral margins of the male pleopod 1, and the topology of the pereiopods, which are ventro-laterally inserted and directed outwards in Lepidocharon gen. n. and dorso-laterally inserted and directed ventrally in Microcharon. Lepidocharon shares with the genus Janinella the morphology of the tergites and the reduced lacinia mobilis of the left mandible, but differs significantly from Janinella in having a well-developed antennal scale, very reduced coxal plates also in females bearing oostegites, the general morphology and spatial arrangement of the stylet-guiding groove of male pleopod 1 and the possession of a 6-segmented antennula. The family Microparasellidae is redefined as monotypic, the only genus being Microparasellus Karaman, 1933.
Isopoda , Microparasellidae , Lepidocharontidae new family, Lepidocharon new genus, coral reef, Australia
The marine isopod fauna of Queensland is diverse with 343 recorded species. Of that total only 22 species (6.4%) are Asellota, the suborder clearly being under-documented for the region as worldwide (
Describing what proved to be a new genus of Asellota led to a reappraisal of the Microparasellidae, and to the conclusion that the family should be split, with the new genus being placed in the Lepidocharontidae Galassi & Bruce, fam. n. A further three species of Lepidocharontidae have been collected from the Great Barrier Reef, including Heron Island in the south, these being undescribed species of Microcharon and Lepidocharon, respectively, and a distinctive new species of uncertain generic identity, yielding a total of at least five species in three genera for the GBR. The presence of this number of species suggests that with appropriate collecting in suitable habitats this family may be more diverse in marine habitats that previously believed.
Collection methods have recently been described by
Sampling was carried out under GBRMPA Permit G08-27858.1 and General Fisheries Permit (QLD DPI) 95152.
The type material is deposited in the Museum of Tropical Queensland, Australia (MTQ).
Male. Body dorsally flat, slender, ~4–10× long as wide, without chromatophores; somites all subsimilar in width, somites sub-rectangular or trapezoidal, lateral margins of head and pereionites sub-parallel. Pleon of one segment, with free lateral margins. Head with weak or absent rostrum, without pseudorostrum. Eyes absent. Antennula with maximally 4 flagellar articles. Antenna flagellum longer than podomeres. Antennal scale (rudimentary exopod) present, even if more or less developed among genera. Mandible incisor with 2 to 8 cusps; molar process subconical, without grinding surface, with apical unequal smooth and pinnate setae; spine row and lacinia mobilispresent, the latter only on left mandible. Maxilliped slender, covering entire mouthpart field, endite distal margin narrowly rounded; epipod slender, quadrate or distally acute; palp composed of 5 articles; 2 stiff pectinate setae always present on maxilliped distal article. Pereiopods 1–7 subsimilar, always without subchela; all pereiopods with 2 dactylar claws; pereiopods articulating dorso-laterally or laterally, and projecting ventrally (in Janinella and Microcharon) or outwards (in Lepidocharon). Penial processes with openings coalescent and medial. Male pleopods 1 and 2 not operculate; male pleopod 1 distally rounded or subtruncate, with or without acute distolateral lobes; proximal part of the pleopod with or without scale-like elements on postero-lateral margins; stylet-guiding grooves running parallel to the lateral free distal margin of pleopod 1 and folded by a hyaline lamella (transversal and unfolded in Janinella); pleopod 3 endopod with 3 plumose setae (marine taxa) or without plumose setae (freshwater taxa), exopod slender. Pleopod 4 globular, pleopod 5 absent. Uropods biramous, ventrally inserted on pleotelson, protopod large, c. 0.5–1.3 as long as pleotelson; protopod length/width ratio c. 2.5–4.5; rami slender with exopod articulating anteriorly to endopod. Anus terminal, not covered by pleopods. Anus outside pleopodal chamber, between bases of uropodal protopods.
Female. Operculum (pleopod 2) from sub-quadrate (as long as wide) to more than 2 times longer than wide, with free distal margin deeply incised medially, faintly incised, or without medial incision, armed with 4 or 2 setae, or unarmed.
Microcharon Karaman, 1934; Janinella Albuquerque, Boulanouar & Coineau, 2014; Lepidocharon Galassi & Bruce, gen. n.
Angeliera Chappuis & Delamare Deboutteville, 1952.
The description of the new genus Lepidocharon Galassi & Bruce, gen. n. led to a re-appraisal of the taxonomic status of the Microparasellidae and its constituent genera. We conclude that the Microparasellidae is a mono-generic family supported by a prominent acute or narrowly rounded rostrum, the antennal flagellum shorter than podomeres, all somites with straight lateral margins that also have scales, an indisputable ventral position of the pereiopods, the unique uniramous and short uropods (see Appendix
Within the family Lepidocharontidae there is great uniformity of the diagnostic characters among all the genera. The genus Angeliera has been placed incertae sedis in the Asellota on the basis of marked differences in several morphological features that set this genus far away the basic body plan observed in Lepidocharontidae.
The largest genus in the Lepidocharontidae is Microcharon with 77 species, both marine and freshwater. Many species lack full descriptions, and there are inconsistencies in the distribution of certain characters within the genus. A dorsal view of the head is not routinely figured; when figured, it can be seen that some species do have a rostral point or rostrum, while others have the anterior margin of the head weakly concave.
The generic name Microcharon is unavailable under the
The family name Microparasellidae was first introduced by
Conversely, we provisionally maintain current and common usage of the names Microparasellus and Microcharon, and this is discussed in more detail together with a new diagnosis for the Microparasellidae (see Appendix
1 | Uropods uniramous, short; antennal podomeres longer than flagellum, scale missing; head with prominent rostrum | Microparasellidae (Microparasellus) |
– | Uropods biramous, large; antennal podomeres shorter than flagellum; rostrum small or absent | 2 |
2 | Pereionites 1–7 cylindrical, free pleonite as wide as pereionite 7 | Microcharon |
– | Pereionites 1–7 dorsally flat and trapezoidal, except pereionite 4; free pleonite narrower than pereionite 7 | 4 |
3 | Male pleopod 1 with transverse stylet-guiding grooves, unfolded; proximal postero-lateral margins of male pleopod 1 without scale-like elements | Janinella |
– | Male pleopod 1 with distal stylet-guiding grooves parallel to the lateral margins, folded by hyaline lamella posteriorly; proximal lateral margins of male pleopod 1 armed with scale-like elements | Lepidocharon gen. n. |
Lepidocharon priapus Galassi & Bruce, sp. n.; here designated.
Lepidocharon lizardensis Galassi & Bruce, sp. n.
Male. Body slender, 8.5–9.7 as long as wide. Free pleonite narrower than pereionites and pleotelson, visible in dorsal view. Cephalon medio-frontal margin not produced, anterior margin straight, rostrum absent. Pereionites 1–3 anteriorly widest, with distinct anterolateral angle, pereionite 4 sub-rectangular, pereionites 5–7 posteriorly widest, lateral margin with distinct posterolateral angle. Pereionites dorsally ornamented by paired setae. Cuticle with small semicircular thickening present or absent both dorsally and ventrally. Antennula 6-segmented; long aesthetascs on articles 5 and 6, long brush seta on article 2 extending to tip of article 6. Antenna with 6 podomeres; article 3 with long blade-like or candle flame-like scale, reaching article 5; lateral margin with 2 setae; flagellum with 8–12 articles.
Mandible palp article 1 unarmed, article 2 with 2 stiff spinulose setae, article 3 with 5 stiff spinulose setae. Right mandible: incisor with 6 to 9 cusps; lacinia mobilis absent; molar process conical, with 3 setae. Left mandible: incisor with 2 to 3 cusps; lacinia mobilis present and produced in 2 cusps; molar process conical, with 3 setae. Maxillula: mesial lobe slender and tapering at distal part, bearing 1 short apical seta accompanied by subapical shorter setae and lateral thin and short setae. Lateral lobe sub-rectangular in shape. Apical setation composed by a variable number of setae. Maxilla: mesial ramus with 8–9 setae; 1 apical comb-like seta, strong, unipinnate and ornamented with fine regularly-spaced setules parallel to one another. Lateral rami close-set, each bearing 4 slender and simple setae, respectively. Maxilliped palp wider than endite, mesial margin of articles 2 and 3 expanded. Pereiopods all subequal in length, all subsimilar in size and general morphology; all with 2 dactylar claws; pereiopod 1 dactylar claws subequal; pereiopods 2–7 superior claw slender (3.8–4.3× basal width), inferior claw robust (2.3–2.6× basal width). Coxae rudimentary reduced to small sclerites, not discernible in dorsal view, coalescent to body wall of the sternites, located on the anterior margin of the concavity which houses the propodus, and apparently not articulated to the sternites.
Pleotelson 1.3–1.7 as long as wide, 1.7–1.9 as long as pereionite 7. Penial papillae opening on postero-medial margin of sternite 7. Pleopod 1 rami proximally fused; proximolateral margins with cuticular imbricate scales on posterior side; distolateral margins convex; stylet-guiding groove represented by a folded hyaline lamella running sub-parallel to free lateral margin of rami; pleopod 1 transverse stylet-guiding grooves absent, unlike Janinella. Pleopod 2 stylet long and slender, of variable length. Pleopod 3 endopod bearing 3 distal plumose setae; exopod elongate, lateral margin with thin setae, article 2 with 1 subapical seta; pleopod 4 rudimentary, ovoid, uniramous. Uropodal protopod as long as the pleotelson, slender (length/width ratio: 4.3), not sexually dimorphic, with long and slender exopod and endopod.
Female. As for the male, except for sexual characters. Operculum (pleopod 2) longer than broad, with surface smooth or with semicircular thickening, with distal margin faintly incised, and 4 apical setae.
The generic name is derived from the ancient Greek name λεπις, λεπιδος meaning “scale”, which refers to the unique rim of scale-like elements bordering the proximal part of the first male pleopod on the posterior side, combined with the mythological name Charon, Charontis referring to the Ferryman of Hades. Gender: masculine.
Lepidocharon gen. n. is most similar to the genus Microcharon, the two genera sharing the following characters: well-developed uropods with slender endopod and exopod; pereiopodal coxal plates not discernible in dorsal view, small, incorporated to the sternite body wall; male pleopod 1 with similar general organization, the distal lateral lobe with a folded hyaline lamella (stylet-guiding groove) running almost parallel to the lateral margins of the pleopod (this orientation and structure of the stylet-guiding groove appears different from that of Janinella, where a transversal and oblique groove hosts the stylet of the male pleopod 2 which seems not to be enveloped by a hyaline lamella); pleopod 2 identical in the development of both exopod and endopod, the latter ending with a stylet of different length depending on the species; penial papillae small and located at the posteromesial margin of sternite 7; female operculum as long as the pleotelson, faintly incised, bearing 4 apical setae (a condition shared by some predominantly marine, and rarely freshwater, Microcharon species).
Lepidocharon gen. n. shares with Janinella the morphology of the tergites, the first three pereionites with antero-lateral protrusions, the fourth sub-rectangular in shape, and the last three postero-laterally protruded, together with the lateral insertion of pereiopods 1–7 oriented outwards (vs. ventrally in Janinella, see
Lepidocharon gen. n. differs from all the other lepidocharontid genera by the combination of the following morphological characters: (1) the unique presence of scale-like elements bordering the postero-lateral margins of the proximal part of the male pleopod 1; (2) pereionites with coxal plates hardly discernible, small, and incorporated into the sternite body wall, not visible in dorsal and lateral views; (3) the long and slender pereiopods that are inserted laterally and directed outwards; (4) pereionites that are not cylindrical, except pereionite 4 (vs. cylindrical in Microcharon); (5) the presence of elongate antennal scale (vs. rudimentary in Janinella, and generally reduced in Microcharon); (6) the mandible incisor with up to 9 cusps (lower number of cusps in Microcharon and Janinella).
Microcharon galapagoensis Coineau & Schmidt, 1979 is closer to Lepidocharon gen. n., especially in the general body morphology, pereionite shape, slender pereiopods directed outwards, a long antennal scale, and slender and elongate uropods; however the unique scale-like elements of the male pleopod 1 of Lepidocharon are not present in this species. These scales are not easily detected under optical microscopy (if not at 100× magnification), nor were they seen using SEM because located on the posterior side of the pleopod.
The mid-section of the male pleopod 1 shows lateral margins expanded in both species of Lepidocharon gen. n. In contrast to the new genus, M. galapagoensis shows a 5-segmented antennula.
Holotype here designated. Adult ♂ (1.3 mm) completely dissected and mounted in polyvinyl lactophenol on one slide (MTQ W28329), 17 February 2009, coll. N.L. Bruce and M. Błażewicz-Paszkowycz.
Type-locality: Australia, Great Barrier Reef, off Coconut Beach, Lizard Island, reef front, sand adjacent to bommies, 4 m, stn LIZ 09–09A, 14.68441°S, 145.47197°E.
Paratypes. 2 ♂♂ (0.9, 1.1 mm), 2 ♀♀ (1.1, 1.4 mm), Great Barrier Reef, Australia, completely dissected and mounted in polyvinyl lactophenol; 2 ♂♂, 1 ♀, same data as holotype preserved in alcohol; 1 ♂, 1 ♀ mounted on SEM stubs (all MTQ W30933).
The epithet is derived from the god Priapus (Πρίαπος) of the Greek mythology. He was considered the protector of livestock, fruit plants, gardens and male genitalia. He was famous for his largely phallic character and the specific names is here referring to the extraordinary length of the stylet of male pleopod 2.
Body length, measured from tip of cephalon to end of pleotelson, from 0.9 to 1.3 mm (n=6). Body 7.0–8.0 times longer than wide, dorsally flat. Cephalon (Fig.
SEM micrographs of Lepidocharon priapus gen. n., sp. n. ♂ paratype. A antennula, sixth segment arrowed B maxillipeds, general view C maxilliped endite and rudimentary sympod (?) arrowed D coxal plates of pereionites 5–7, ventral view E pereiopod 1, detail of dactylus and reduced sclerite F pereiopod 7, detail of dactylus and articulated sclerite G pereiopods, lateral view H rudimentary coxal plates, ventral view, arrowed.
Paragnaths (Fig.
Antennula (Fig.
Antenna (Fig.
Mandible palp (Fig.
Maxillula (Fig.
Maxilla (Fig.
Maxilliped (Figs
Pereiopod 1 (Figs
Pleonite 0.29 as long as and 0.84 as wide as pereionite 7 (Figs
SEM micrographs of Lepidocharon priapus gen. n., sp. n. A, B, C ♂ paratype. A general view of free pleonite (ventral view), and pleopod 1 and 2 B penial papillae with ventral groove (arrowed) C pleopod 1, detail of the anterior proximal opening to the mesial channel D ♀ paratype, operculum.
Penial papillae, as for the genus, coalescent and located at the middle of the free posterodistal margin of pereionite 7, with undulated free outer margins and a medial channel (Fig.
Pleotelson 1.37 as long as maximal width (Figs
Male pleopods 1 elongate and slender, fused proximally, with sperm tube medially with an anterior opening ornamented by small spines (Figs
Male pleopod 2 (Fig.
Pleopod 3 (Fig.
Pleopod 4 (Fig.
Uropods long and slender (Fig.
Body length generally similar to male (Fig.
Detailed comparison between the two species is given in the remarks for Lepidocharon lizardensis sp. n.
Holotype here designated. Adult ♂ (1.1 mm), completely dissected and mounted in polyvinyl lactophenol on one slide, 17 February 2009; coll. N.L. Bruce and M. Błażewicz-Paszkowycz (MTQ W28330).
Type-locality: Australia, Lizard Island, off Coconut Beach, 14.68441°S, 145.47197°E, reef front, sand adjacent to bommies, 4 m, stn Liz 09-09A.
Paratypes. 1 ♂ (0.9 mm), 1 ♀ (1.2 mm) completely dissected and mounted in polyvinyl lactophenol, same data as holotype (all MTQ W28331).
The epithet lizardensis is after the type locality, Lizard Island, northern Great Barrier Reef, Queensland.
Body length measured from tip of cephalon to end of pleotelson 0.9–1.1 mm (n = 2). Body dorso-ventrally flattened, stocky, small-sized, about 6.5 times longer than wide (Figs
Pereionites 1–7 subequal in width (Fig.
Paragnaths (Fig.
Antennula (Fig.
Antenna (Fig.
Mandible. Palp (Fig.
Maxillula (Fig.
Maxilla (Fig.
Maxilliped (Fig.
Pereiopod 1 (Fig.
Pleonite length 0.27 pereionite 7 length, width 0.86 pereionite 7 width (Figs
Pleotelson longer than wide (Fig.
Male pleopods 1 (Fig.
Male pleopod 2 (Fig.
Pleopod 3 (Fig.
Pleopod 4 (Fig.
Uropods unknown.
Body length approximately as in male. Body length measured from tip of cephalon to end of pleotelson 1.2 mm. No sexual dimorphism observed in body morphology, cephalic appendages and pereiopods. Female operculum (pleopod 2) elongate (Fig.
Lepidocharon priapus and L. lizardensis differ from each other in several characters: 1) the morphology of the antennal scale (blade-knife shaped in L. priapusvs. candle-flame shaped in L. lizardensis); 2) the slender body with different degree of protrusion of the pereionites 1–3 and 5–7 (markedly protruded in L. priapus vs. stouter and less protruded in L. lizardensis); 3) the different shape of the male pleopod 1 (with strongly protruded and pointed apical lobes and sclerotized hyaline lamella in L. priapus vs. sub-rounded and undulated apical lobes and a tiny hyaline lamella in L. lizardensis); 4) the setal complement of the male pleopod 1 (7 in L. priapus vs. 6 in L. lizardensis); 5) the morphology of the male pleopod 2 (sub-rectangular and long protopod with endopodal stylet extraordinarily long, reaching the tips of the uropods in L. priapus vs. ovoidal protopod with a short stylet, not reaching the distal part of the protopod in L. lizardensis); 6) female operculum sub-rectangular in shape, with straight lateral margins in L. priapus vs. oval, with convex lateral margins in L. lizardensis); 7) a rounded pleotelson in L. priapus (sub-truncate in L. lizardensis), and 8) body surface with visible semicircular thickening in L. priapus (smooth in L. lizardensis).
The family Microparasellidae has been provisionally assigned to the asellote superfamily Janiroidea Sars, 1897. Its monophyletic status was debated since the interpretation of most character states in microparasellids is still doubtful, if not questionable (
Similar arguments had been anticipated by
Among the genera assigned to the former family Microparasellidae, the genus Angeliera uniquely has a 7-segmented antennula, antennal scale absent; the maxilliped palp 4-segmented, without apical stiff pectinate setae; most pereiopods (variable among species of the genus) with three claws; pereiopods 1–4 and 6–7 subsimilar, with pereiopod 5 sexually dimorphic, in the male being subchelate and stout, with carpus transformed; male pleopod 1 is short and sub-quadrate and the penial processes are paired, not coalescent, arising near the base of pereiopod 7 (i.e. laterally) (see for details
Angeliera lacks the molar process, the most derived condition found in the Janiroidea. Nevertheless, similar reductions of the molar process occur in the janiroid family Katianiridae Svavarsson, 1987, where it varies from being reduced to a single spine in Katianira acarina (Menzies, 1962) (
A single free pleonite is shared by the Janiridae, the Microparasellidae and the Lepidocharontidae fam. n., as well as by other phylogenetically distant families (e.g. Thambematidae Stebbing, 1912; Paramunnidae Vanhöffen, 1914; Urstylidae Riehl, Wilson & Malyutina, 2014). Members of the genus Microcharon, as observed in Microcharon reginae Dole & Coineau, 1987, possess a large free pleonite (Fig.
Stereomicroscope images of the habitus of A Lepidocharon priapus gen. n., sp. n. ♀ paratype B Lepidocharon lizardensis gen. n., sp. n. ♀ paratype C Microcharon reginae Dole and Coineau, 1987 ♀ topotype, showing the different morphology of the pereionites, the topology of pereiopods, and the different degree of development of the single pleonite free (scale bars: 0.5 mm).
The single free pleonite is even more reduced in the janirid Microjaera anisopoda Bocquet & Lévi, 1955, as highlighted by
The relatively large size of the free pleonite in Microparasellidae (Microparasellus) has been considered a distinctive trait of the family by
The antennula in the Lepidocharontidae is short, not sexually dimorphic, and composed of a maximum of 6 articles, this condition being derived in the Janiroidea. The segmental pattern of the antennula ranges from 6 to 5 in Microcharon, where the 5-segmented condition is shared by almost all the marine species of the genus, and the 6-segmented condition by fresh groundwater species, which likely retained the 6-segmented antennula of the ancient marine ancestor, in the more conservative and stable groundwater environment, according to
The antennal exopod (scale) is present in all the Lepidocharontidae genera. In general, the exopod is rudimentary in the Janiridae, and in members assigned to the Lepidocharontidae is ovoid and small, as in Microcharon, the most reduced condition being found in Janinella (
The molar process of the mandible is always conical and reduced in the lepidocharontid genera Microcharon, Janinella and Lepidocharon.
In Lepidocharon the mandibular palp is inserted on short cuticular projection, as also observed in Janiropsis Sars, 1897 and Janaira Moreira & Pires, 1977 by
The distal article of the mandibular palp bears from 5 to 3 robust setae, the latter status shared by the most derived groundwater species of Microcharon. The possession of 5 setae is shared by a few marine Microcharon species, when described, and the stygobiotic Microcharon acherontis Chappuis, 1942 is figured with 6 setae, a state not found in any other member of the genus.
In the incertae sedis genus Angeliera, the terminal segment of the mandibular palp has been described and figured without any seta or stiff setae, and some descriptions refer to the original description of A. phreaticola Chappuis & Delamare Deboutteville, 1952 for which few details are available (Chappuis and Delamare 1952,
The maxilliped is 5-segmented in Microcharon, Janinella and Lepidocharon.
The genus Angeliera placed as incertae sedis in the Lepidocharontidae has a 4-segmented palp as other janiroid families, e.g., Katianira and Natalianira Kensley, 1984 (Katianiridae), raising questions about purported affinities between katianirids and the genus Angeliera (see
The maxilliped endite has the same shape, recurrent in all members of the family, except for the genus Angeliera, where its distal part is swollen and ending with sinuous setae.
The body morphology is generally similar among members of the family Lepidocharontidae: pereional somites are rectangular in Microcharon, with a well-developed free pleonite, shorter and as wide as pereionite 7. Janinella and Lepidocharon have the anterior three pereionites markedly protruded anteriorly, the fourth almost rectangular in shape, and the last three pereionites protruded posteriorly. We assume that the morphology of the pereionites, together with the position of the pereiopods are related to a different way of locomotion of the species.
The pereiopodal dactylus has two unguli, showing a general tendency to be stout and subequal in length in marine species, and slender with the inner claw about ½ the length of the lateral claw in freshwater species.
The morphology and armature of the female operculum differs within the family. A character considered diagnostic for species distinction within the family is the number of apical setae bordering the female operculum. In Microcharon the highest variation occurs: from long, more than two times longer than wide, faintly incised female operculum in marine and freshwater species, bearing 4 or 2 apical setae (
The pleopod 3 has the distal article of the endopod either with 3 plumose setae or completely unarmed. The marine species always have these setae and they are consistently absent in all groundwater species.
Angeliera shows the most striking derived character states, suggesting a divergent position in relation to the remaining members of the family. Janinella shares with Angeliera the topology of the outwardly directed stylet-guiding groove of the male pleopod 1 on both pleopodal rami, a character that may be phylogenetically informative, and likely primitive in the family because it is shared by members of the closely related family Janiridae (e.g. Jaera Forsman, 1949, Iais, Ectias Richardson, 1906). Microcharon and Lepidocharon share stylet-guiding grooves folded by a hyaline lamella that run almost parallel to the free lateral margins of the distal parts of male pleopod 1, a likely derived organization and topology of the stylet-guiding groove if compared to that of Janinella. The stylet guiding-groove is in this case represented by a hyaline lamella, which holds the stylet of the pleopod 2 in a precise position. In the first case, as in Jaera (
The penial papillae are differently organized in Angeliera: they are tubules which start from the insertion of the pereiopods on sternite 7 and converge at the midline of the sternite of the pereionite 7, maintaining their openings separate and not coalescent (
The description of Lepidocharon Galassi & Bruce, gen. n. and the establishment of the Lepidocharontidae Galassi & Bruce, fam. n. shed new light on the diversity of the morphological body plans observable in the former family Microparasellidae, allowing for a better understanding of the phylogenetic relationships of the Lepidocharontidae fam. n. and the Microparasellidae, a family placed as incertae sedis within the superfamily Janiroidea Sars, 1897 (see
The small body size, soft cuticle, possession of the single reduced pleonite with laterally free margins, the simplification of the general structure and armature of the mandibles, the low diversification of setae morphology, the absence of pleopod 5, could be interpreted all together as derived character states, which may conversely be the result of homoplasy for adaptive convergence due to the interstitial life, or to life in microhabitats with reduced living space (
In addition, the lack of morphological data in both historic and some recent contributions does not help in reconstructing an evolutionary scenario. Nonetheless, despite convergence, phylogenetic characters can be identified, and a greater degree of character resolution in family and genera definitions developed. Even if comprehensive data for all species are not available, it is evident that the current generic composition of the Microparasellidae could not be maintained. The separation of the Lepidocharontidae fam. n. and its constituent genera resolves the former paraphyletic family into two monophyletic families, which in future should enable a more clear understanding of the relationships of the two families to the Janiridae to be developed.
The authors gratefully acknowledge the Australian Biological Resources Study for providing support to NLB for the early part of this study (ABRS grant 2009–30). Material from Lizard Island was collected under the auspices of the CReefs project organized by the Australian Institute of Marine Science (AIMS). The CReefs Australia Project is generously sponsored by BHP Billiton in partnership with The Great Barrier Reef Foundation, the Australian Institute of Marine Science and the Alfred P. Sloan Foundation; CReefs is a field program of the Census of Marine Life. We thank Julian Caley and Shawn Smith (AIMS) for their excellent organization and field support; NLB thanks Magdalena M. Błażewicz-Paszkowycz for her excellent team spirit while collecting at Lizard Island. Sandro Minelli (member of the ICZN Commission) is greatly acknowledged for the useful suggestions given to DMPG. Nicole Coineau kindly supported DMPG in improving the original manuscript. An earlier version of the manuscript greatly benefited from the comments of three reviewers.
The family name Microparasellidae was first formally proposed by
Synonymy: Karaman 1934: 44.—Wolff 1962: 35.—Coineau 1986: 465.—Kensley and Schotte 1989: 90.—Wilson and Wägele 1994: 720.
Body slender, 4.0–6.0× long as wide, somites all subsimilar in width, head with acute or narrowly rounded rostrum. Lateral margins of head and pereionites convex, with cuticular scales; pleonite 1 laterally free; body without chromatophores. Eyes absent. Antennula flagellum with maximally 4 articles. Antennal scale absent. Antenna flagellum shorter than podomeres. Mandibular molar process distally pointed, without grinding surface, with several terminal setae. Pereiopods 1–7 not chelate or subchelate, almost identical in shape; all pereiopods with 2 dactylar claws. Pereiopods articulated latero-ventrally. Penial processes present, coalescent, with single medial opening on posterior margin of pereionite 7. Pleopods 1 and 2 not operculate in males; female pleopod 2 operculate; pleopod 3 endopod unarmed, exopod slender. Uropods uniramous, minute, stub-like, insertion ventro-terminal; single ramus, shorter than protopod. Anus terminal, not covered by pleopods.
The family Microparasellidae was ambiguously established by
Microparasellus Karaman, 1933: 17; 1934: 44.—Wilson and Wägele 1994: 725.
Duslenia Lévi, 1950: 42.
Microparasellus puteanus Karaman, 1933. Type locality: Skopje, Macedonia.
Microparasellus libanicus Chappuis & Delamare Deboutteville, 1954, Lebanon; Microparasellus aloufi Coineau, 1968, Lebanon; Microparasellus hellenicus Argano & Pesce, 1979, Greece.
The generic name Microparasellus established by
The Microparasellidae can be regarded as monophyletic as here defined. While we do not enter a discussion into monophyly or otherwise of the Janiroidea, it is apparent, as shown by
Species of the family are known from North Africa and eastern Europe only.