taxonID	type	description	language	source
03AB87850463FFEB0770FDD0FD1EFA47.taxon	materials_examined	Type genus. Haplomunna Richardson, 1908. Diagnosis (modified from Wilson 1976): Head without eyes, anteriorly rounded, dorsal surface sloping anteriorly, lacking anterior margin. Body with spines, dorsally vaulted, posterior pereonites axially compressed and reduced. Female spermathecal duct emerging anterodorsally near margin of pereonite 5. Anus covered by opercular pleopods. Antennula inserting dorsally on head, sexually dimorphic, with many flagellar articles and aesthetascs in male, and fewer and thinner articles in female. Antenna inserting anterodorsally, basal articles robust, article 3 without scale. Mandible palp (when present) elongate, longer than body of mandible, article 3 elongate with elongate setae only on distal tip. Pereopod I with major subchelate hinge between carpus and propodus; carpus palm (ventral margin) robust, with elongate robust setae. Pereopods II – VI and VII, if present, longer than body. Male pleopod II protopod strongly tapering and narrow distally; exopod lacking setose posterior projection. Pleopod III exopod biarticulate, narrower and longer than endopod, without plumose setae. Uropods emerging at pleotelson ventral margin; uniramous; tiny, not extending beyond terminal margin of pleotelson. Relationships: The family Haplomunnidae was argued by Wilson (1976) to be the sister group of the Dendrotiidae, and not closely related to Munnidae, Santiidae or Paramunnidae. The primary similarities between the former two families include general body spinosity, vaulted thoracic region, compact anteriorly­sloping emarginate head capsule with antennae and antennulae placed on anterodorsal surface, and elongate pereopods II – VII that exceed the body length. While not fully consistent among species in both families, the ischium and merus of the walking legs are often subequal and shorter than the basis. Notably, the opposite situation occurs among the Munnidae, where the basis is often shorter than the ischium. Although Acanthomunna Beddard, 1884 is somewhat similar in body form to Haplomunna, male dendrotiids have a setose posteriorly­directed appendage on exopod of pleopod II, an apomorphy of the family. Tiny marginal uropods are an apomorphy of the Haplomunnidae, although reductions of the uropods are common among other deep­sea isopod taxa. Some, but not all, members of both families share the heterochronic non­expression of the last pereonite. This oddity also occurs among other deep­sea families (e. g., Munnopsidae, Ischnomesidae), so its phylogenetic significance is limited to those clades that reach adulthood without the last pereonite. Regarding higher­level relationships, Wägele (1989), partially following Wolff (1962), suggested that the asellotan families Munnidae, Santiidae (as Pleurocopidae), Dendrotiidae and Haplomunnidae were related in a group referred to as ‘ munnoiden’, or as having a ‘ munnoid habitus’. The sister group relationship between the Dendrotiidae and the Haplomunnidae, as discussed above, was not in dispute, but the principal difficulty with Wägele’s (1989) hypothesis lies in the underlying assumption of similarity of body form (Wilson 1996). In Wägele’s (1989: 67) concept, the munnoid habitus includes the following features: an approximately oval body outline, in which the middle pereonites are somewhat broader than the anterior or posterior segments, head broader than long, eyes laterally projecting and often clearly stalked, pereonites 5 to 7 not only narrower but also shorter than the anterior pereonites, pleotelson dorsally convex, and at its origin somewhat narrower, distally rounded and often spindle­like. Other than the stalked eyes, most of these features are seen throughout the Janiroidea, especially among the deep­sea families. Acanthomunna is the only genus with eyes among the Dendrotiidae or the Haplomunnidae, but only has rounded lateral projections, somewhat similar to those seen among some Janiridae and clearly unlike the stalked eyes of the presumptive ‘ munnoiden’ clade. Although not fully relevant here, the form of the stalked eyes (where present) varies consistently between the eyed families: short, thick and generally with many ocelli among the Munnidae and Santiidae, elongate and often with few ocelli among the Paramunnidae and extremely elongate and thin, with few ocelli in Pleurocope, which belongs to its own separate family, Pleurocopidae. The ‘ munnoiden’ are unlikely to be a cohesive phylogenetic group because different taxa have different positions for the spermathecal duct and configurations of pereopod I (Wilson 1987; personal observation). Additionally, antennae in Paramunnidae, Munnidae and Santiidae, and Pleurocopidae all insert anteroventrally, but with differing positions among the three groups; Haplomunnidae and Dendrotiidae have antennae inserting anterodorsally, often on projections of varying sizes. Other features, such as the above­mentioned comparative lengths of the pereopodal podomeres, also differ considerably among these families. Clarification of the haplomunnid classification will require detailed morphological study and phylogenetic analysis of all families of Janiroidea. Distribution: Although Kussakin (1988), in his useful compendium of the isopods of the Northern Hemisphere, characterised the Haplomunnidae as an equatorial and Southern Hemisphere group and only listed the equatorial Atlantic species Thylakogaster peterpauli Wilson & Hessler (1974), recent records (Table 1, Figure 1) indicate that this family is much more widespread than previously suspected, and clearly not limited to equatorial regions. Species of Haplomunna are known from abyssal and hadal waters of the North Pacific, off southern and Baja California (Wilson 1976; Beaulieu 2001 a, 2001 b) and the Japan Trench (Gamô 1983) from depths of 3880 to 6450 m. Thylakogaster has the broadest distribution from depths of 1135 m to 5223 m, including the equatorial Atlantic, Argentine Basin, north of Surinam, Bermuda slope, off Ireland, in the Rockall Trough (Harrison 1988) and in the east Pacific (Thistle & Wilson 1987). New records include the Lucky Strike vent field on the Mid­Atlantic Ridge, and the east Pacific Ocean. Abyssaranea has been found at depths from 3459 to 4438 m in the equatorial Atlantic and East Pacific, as well as a new record from a Woods Hole Oceanographic Institution deep­benthic station in the northwestern Atlantic. This latter record is the first non­equatorial record for this genus. Munella has most records in the bathyal deep sea (950 – 2076 m) including the Bay of Biscay, the Bermuda Slope and north of Surinam in the Atlantic. Unconfirmed records of Munella in the central Mediterranean are from depths of 100 – 200 m (Bay of Napoli; Lo Bianco 1903). Munella also occurs at abyssal depths in the east Pacific (4797 m, Thistle & Wilson 1996). New records of Munella include Lucky Strike vent field on the Mid­Atlantic Ridge (1685 m, TTR­ 10), the eastern Atlantic near the African coast (1175 – 1283 m, BALGIM stations), and the Gulf of Mexico (3732 m., Rowe 2003). A new species and new genus of Haplomunnidae collected by Poore et al. (1994) occurs in southeastern Australia at bathyal depths (J. Just, personal communication). Habitat and Ecology: All Haplomunnidae have long­legged ambulatory morphologies, and are unlikely to be burrowers. Because they are primarily epibenthic and exposed to erosion, they are not likely to be found at high energy sites. This family has not been recorded from the HEBBLE (high energy benthic boundary layer experiment) sites (Thistle & Wilson 1987, 1996), and haplomunnid species are a reasonably consistent member of benthic assemblages at quiescent sites. In the Clipperton­Clarion Fracture Zone region of the East Pacific, a region characterised by low abyssal hills, pelagic clay and manganese nodules, Thylakogaster has been found at the ECHO 1 site (Thistle & Wilson 1987) and Munella at the PRA site (Wilson 1990; Thistle & Wilson 1996). The recorded substrates for haplomunnids range from typical deep­sea sediments to rocky outcrops, manganese nodules and carbonate crusts. Several Atlantic records are from continental margins in muddy sediments (Bonnier 1896), sand and volcanic gravel (Lo Bianco 1903), mixed Globigerina and pteropod oozes (Bermuda slope; Wilson & Hessler 1974). We now report them from hydrothermal vents (Lucky Strike) including a high temperature, diffuse venting area. In this case, the Thylakogaster specimens were collected on volcanic rocks (hyaloclastic breccia and pillow lavas) in the vicinity of active venting areas and in colonisation trays deployed near black smokers in the SE part of the field, while Munella specimens were collected on fragments of inactive chimneys in the NW part of the field. Biogenic structures appear to be favoured, because Haplomunna sp. was found highly aggregated on Hyalonema sponge stalks (Beaulieu 2001 a, 2001 b). Like most deep­sea isopods, the trophic habit of Haplomunnidae species appears to be mostly detritivory. Haplomunna sp., found on the basal and middle portions of hexactinellid sponge stalks, may be feeding upon particles accumulating in pockets (Beaulieu 2001 a). Thylakogaster species have elongate and setose pereopods II – III that may be used to search food in the sediments or catching suspended particles. These legs on preserved specimens are generally held with the carpus and propodus curving under the mouth field and the first pereopods, like a basket.	en	Cunha, Marina R., Wilson, George D. F. (2003): Haplomunnidae (Crustacea: Isopoda) reviewed, with a description of an intact specimen of Thylakogaster Wilson & Hessler, 1974. Zootaxa 326: 1-16, DOI: 10.11646/zootaxa.326.1.1
03AB87850460FFE40770FA56FAA7F9E8.taxon	materials_examined	Type species. Munna caeca Richardson, 1905, by monotypy. Species included: Haplomunna caeca (Richardson, 1905); Haplomunna hubbsi Wilson & Hessler, 1976; Haplomunna japonica (Gamô, 1983). Diagnosis (modified from Wilson 1976): Head rounded in dorsal view; depth and width less than pereonite 1, with distinct frons­clypeal ridge, antennae on low projection. Body ovate in dorsal view, vaulted in lateral view; surfaces and margins spinose, with two distinctly different sizes of spines: large robust spines, often on raised portions of the cuticle, and small thin spines (‘ stiff hairs’ in Richardson 1905), pereonites 5 – 7 with spines only on lateral edges, dorsal surfaces smooth. Pereonites 1 – 4 sculptured, dorsal surfaces often with parallel transverse ridges arching across dorsum; lengths subequal. Female coxal developing oostegites external to sternal surface. Pereonites 5 – 7 dorsomedial axis compressed; lateral portions extending posteriorly beneath elevated pleotelson. Preparatory female with external developing oostegites. Pleotelson longitudinal axis nearly perpendicular to pereonal axis; inflated, width subequal to pereonite 5 width, somewhat bilobed anterodorsally, heart shaped in dorsal view; distal tip smooth, produced ventroposteriorly. Mandible molar process truncate with two distal ridges: distal ridge finely denticulate; proximal ridge with inner row of stout spines, and outer row of long sharp, minutely serrate spines; palp elongate, longer than body of mandible, article 1 longer than articles 2 – 3, article 3 subequal to article 2, with elongate setae in group at distal tip. Maxillipedal palp with segment 2 distinctly broader, segment 3 only slightly broader than segment 1. Pereopod I strong, subchelate; carpus somewhat elongate; distinct rows of many stiff unequally bifid setae on occlusive margins of both carpus and propodus. Pereopods II – VII similar to each other. All pereopods with two dactylar claws. Opercular pleopods with strong spines. Uropod tiny, uniarticulate, directed ventroposteriorly. Remarks: We propose a synonymy of Haplomunna Richardson, 1908 and Aryballurops Gamô, 1983. The species H. japonica differs from the East Pacific species H. caeca and H. hubbsi in having a much more inflated pleotelson, but in all other respects, the three species are similar. Gamô (1983) provides the first description of a male in the Haplomunna clade, details of which we can confirm from specimens of an undescribed species provided by S. Beaulieu (SIO). In particular, the tip of the male pleopod I is rounded and not laterally produced (i. e., not saggitate) as in Thylakogaster, thus having a form more like that of Abyssaranea or Munella. An elongate mandibular palp, another similarity with the latter two genera, and the less modified form of pereopods II – VII clearly separates Haplomunna from Thylakogaster. Nevertheless, Thylakogaster and Haplomunna are similar in several respects, the most conspicuous being the reflexed and inflated pleotelson.	en	Cunha, Marina R., Wilson, George D. F. (2003): Haplomunnidae (Crustacea: Isopoda) reviewed, with a description of an intact specimen of Thylakogaster Wilson & Hessler, 1974. Zootaxa 326: 1-16, DOI: 10.11646/zootaxa.326.1.1
03AB8785046FFFE50770F9C5FD3DFBE0.taxon	materials_examined	Type species. Thylakogaster peterpauli Wilson & Hessler, 1974, by original designation. Species included: Thylakogaster peterpauli Wilson & Hessler, 1974; Thylakogaster majusculus Wilson & Hessler, 1974; Thylakogaster lobotourus Wilson & Hessler, 1974. Diagnosis (modified from Wilson & Hessler 1974): Head laterally angular or curved in dorsal view; with transverse ridge and frons­clypeal furrow; depth subequal or greater than pereonite 1. Body with many thin­walled spines or spinules, including cuticle of dorsolateral sides of head, lateral and posterior sides of pleon, and external surfaces of pleopods 1 and 2; spines also on coxae, lateral margins of pereonites 1 and 2; dorsum of pleon devoid of such spines. Female developing oostegites on coxae internal to sternal surface. Pleotelson strongly inflated, hollow, expanded dorsally and laterally into somewhat bilobed form, reflected anteriorly, lying above pereonites. Mandible without palp; molar process tapering distally into tuft of thin, curved spines. Maxilliped epipodite triangular, small, width less than basis width, length less than half basis length. Pereopods II and III elongate, distinctly longer than head – pereon length; carpus and propodus curving medially, strongly opposing each other, combined lengths exceeding two­thirds total pereopod length, with large curved setae; dactylus short, covered with fine cuticular hairs, with single claw. Pereopods IV – VII longer than pereopods II and III, more slender and straight, with smaller setae and with long, slender claw and second vestigial claw on dactylus. First pleopod copulatory male posterior margin sagittate. Uropods uniarticulate, positioned ventroposteriorly close to tip of pleon.	en	Cunha, Marina R., Wilson, George D. F. (2003): Haplomunnidae (Crustacea: Isopoda) reviewed, with a description of an intact specimen of Thylakogaster Wilson & Hessler, 1974. Zootaxa 326: 1-16, DOI: 10.11646/zootaxa.326.1.1
03AB8785046EFFE10770FBF7FC66FD78.taxon	materials_examined	Type material. — NW Atlantic: Bermuda Slope, southeast of northern tip of the Bermuda Islands, 32 º 19 ’ N, 64 º 32 ’ W, 1135 – 1153 m, stn WHOI 118 a, holotype 1 copulatory male (USNM 141471), paratypes 1 immature female (USNM 141472) and 2 copulatory males (ZMUC). Material examined. — Mid­Atlantic Ridge: Lucky Strike, 37 º 17.36 ’ N, 32 º 16.49 ’ W, 1695 m, Submersible Nautile, colonisation tray, stn PICO SMAC C 6, 6. VII. 1998, D. Desbruyères, 1 female (illustrated herein) head – pereon length 1.46 mm (CENTOB). Same locality, 37 º 17.44 ’ N, 32 º 16.60 ’ W, 1685 m, RV Prof. Logachev, TV­assisted grab, stn TTR 10 AT­ 265 ­ Gr, 10. VIII. 2000, M. R. Cunha, 1 manca IV – V head – pereon length 1.24 mm (DBUA). Same locality, 37 º 17.30 ’ N, 32 º 16.56 ’ W, 1709 m, RV Prof. Logachev, TVassisted grab, stn TTR­ 12 AT­ 436 ­ Gr, 29. VIII. 2002, M. R. Cunha, 1 female head – pereon length 3.07 mm, and 1 anterior fragment. NW Atlantic: Bermuda Slope, southeast of northern tip of the Bermuda Islands, stn WHOI 118 a (type locality), 14 individuals or fragments, most in poor condition (AM P 64368); 32 º 16 ’ N, 64 º 32 ’ W, 2095 – 2223 m, stn WHOI 119, 1 individual, pleotelson only (AM P 64369). Supplementary description: Female (Figs. 2 – 3): Head – pereon length 1.46 mm measured laterally. Antennula with 11 articles, article 1 with 2 penicillate setae, article 2 with 1 penicillate seta, article 4 with 1 penicillate seta, all located distally; article 3 length 1.6 article 2 length; article 4 length 0.28 article 2 length (proportions similar to T. peterpauli and T. majusculus). Antenna length 2.6 head – pereon length; article 5 length 2.5 articles 1 – 4 length altogether; article 6 length 1.7 article 5 length, both articles bearing numerous long setae; flagellum length 0.30 article 6 length, with 10 articles. Pereopod I basis length 2.3 width; ischium with 5 ventral setae, 2 dorsal setae; merus with 3 ventral setae, 1 dorsal seta, 4 medial setae; carpus length approximately 2 width, slightly longer than basis, with 11 – 12 setae on ventral margin, with 5 – 6 short stout setae distal to large central seta, proximal medial surface with 10 setae in two rows; dactylus length approximately 0.85 propodus length. Pereopod II long, length 2 head – pereon length; basis length approximately 3 width; propodus slender and curved, length approximately 15 width; both carpus and propodus with numerous setae arising from short cuticular protuberances, setae largest on occlusive margin on carpus and lateral and medial side of propodus; dactylus length 0.2 propodus length, with 1 claw. Pereopod III shorter than but similar to pereopod II. Pereopods IV – VII less robust and longer than pereopod II, length approximately 1.1 pereopod II length; basis width approximately 0.8 pereopod II width; propodus and carpus approximately two­thirds total pereopod length; propodus slender, length 1.4 carpus length (somewhat longer in pereopod VII); propodus with long robust mediodistal seta at dactylar insertion. Remarks: Although the Lucky Strike specimens are distant from the original Bermuda Slope type locality of T. lobotourus, we found no differences that require the establishment of a new species. These new specimens suffered some bending and twisting of their thin legs, so the whole body illustration (Fig. 2 A) was made by merging several views of the limbs to obtain an approximation of the ‘ natural’ condition. Males and juvenile specimens (mancas or females) of Thylakogaster species have usually one or two pairs of setae on the distal margin of pleotelson (between the uropods), and indentations of the margin; mature females lack these setae or indentations. Our observations on Thylakogaster specimens held at the Australian Museum show the following patterns: T. lobotourus WHOI 118 a (AM P. 64368), female no setae, male two pairs of setae; T. lobotourus Lucky Strike female, one pair of setae and indentations (Fig. 2 D); T. majusculus WHOI 242 a (AM P. 64371), juvenile female no setae, male two pairs of setae and indentations; T. peterpauli WHOI 155 (AM P. 64372), small male several small setae, female no setae, both with abundant cuticular hairs and no indentations; T. n. sp. 1 WHOI 303 (AM P. 64373), manca two pairs of setae. The cuticle of T. peterpauli has fine spinules, especially on the pleotelson, that are missing in T. lobotourus. We have observed several differences between species of Thylakogaster: i) The pereopod I carpus setae distal to large central seta on ventral margin differ: 5 or 6 in T. lobotourus, 7 in T. peterpauli; 7 – 9 in T. majusculus. ii) The antenna of T. lobotourus is longer, with a longer flagellum and smaller setae than in T. peterpauli (antenna ~ 3.1 and 2.5 head – pereon length for T. lobotourus and T. peterpauli, respectively). iii) The antennula of an adult male is slender with fewer aesthetascs in T. peterpauli and more robust in T. lobotourus and T. majusculus. iv) Spines on body and pleotelson differ among the species. Relative to each other, number of spines varies as follows: T. majusculus> T. peterpauli> T. lobotourus. The length of the spines shows a different trend: T. lobotourus> T. majusculus> T. peterpauli. The pleotelson in T. lobotourus has fewer spines (10 – 15 % less for the same area) but longer spines (1.1 times) than in T. peterpauli.	en	Cunha, Marina R., Wilson, George D. F. (2003): Haplomunnidae (Crustacea: Isopoda) reviewed, with a description of an intact specimen of Thylakogaster Wilson & Hessler, 1974. Zootaxa 326: 1-16, DOI: 10.11646/zootaxa.326.1.1
