Two new species and new records of Catapagurus A. Milne-Edwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) from the southwestern Pacific
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Lemaitre, Rafael (2025): Two new species and new records of Catapagurus A. Milne-Edwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) from the southwestern Pacific. Zoosystema 47 (31): 751-764, DOI: 10.5252/zoosystema2025v47a31, URL: https://sciencepress.mnhn.fr/sites/default/files/articles/pdf/zoosystema2025v47a31_pdfa.pdf
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2025 47 31 art. 47 (31) — Published on 3 December 2025 www.zoosystema.com Two new species and new records of Catapagurus A.Milne-Edwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) from the southwestern Pacific Rafael LEMAITRE
Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Catapagurus poupini n. sp., holotype male 3.0 mm, Marquesas, Hiva Oa island, MUSORSTOM 9; stn CP1212, MNHN-IU-2022-4536. Zoosystema est indexé dans / Zoosystema is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Zoosystema est distribué en version électronique par / Zoosystema is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Zoosystema sont référencés par / Articles and nomenclatural novelties published in Zoosystema are referenced by: – ZooBank® (http://zoobank.org) Zoosystema est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris / Zoosystema is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387
751 ZOOSYSTEMA • 2025 • 47 (31) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:8ACF647B-7F16-4114-A86F-A75DE158E189 Lemaitre R. 2025. — Two new species and new records of Catapagurus A. Milne-Edwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) from the southwestern Pacific. Zoosystema 47 (31): 751-764. https://doi.org/10.5252/ zoosystema2025v47a31. http://zoosystema.com/47/31 ABSTRACT Two new species of the genus Catapagurus A. Milne-Edwards, 1880, C. poupini n.sp. and C. thysanodactylus n.sp., from the western pacific are fully described and compared with their morphological closest congeners. The horizontal distribution of two others species is considerably expanded eastward to the Fiji Islands, for C. franklinae McLaughlin, 2004 (previously known from the Coral and Tasman Seas, and New Caledonia); and for C. tanimbarensis McLaughlin, 1997, previously known from the Kai and Tanimbar Islands and Indonesia. RÉSUMÉ Deux nouvelles espèces et nouveaux signalements de Catapagurus A. Milne-Edwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) du sud-ouest du Pacifique. Deux nouvelles espèces du genre Catapagurus A. Milne-Edwards, 1880 du Pacifique occidental sont décrites. La répartition horizontale de deux autres espèces est, par ailleurs, considérablement étendue vers l’est: jusqu’aux îles Fidji pour C. franklinae McLaughlin, 2004, connu auparavant de la mer de Corail, de la mer de Tasman et de Nouvelle-Calédonie ; et pour C. tanimbarensis McLaughlin, 1997, connu auparavant des îles Kei et Tanimbar et d’Indonésie. Rafael LEMAITRE Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, 4210 Silver Hill Road, Suitland, MD 20746 (United States) [email protected] Submitted on 20 November 2024 | Accepted on 30 April 2025 | Published on 3 December 2025 Two new species and new records of Catapagurus A. MilneEdwards, 1880 (Crustacea, Decapoda, Anomura, Paguridae) from the southwestern Pacific MOTS CLÉS Paguridae, Catapagurus, signalements nouveaux, espèces nouvelles. KEY WORDS Paguridae, Catapagurus, new records, new species.
752 ZOOSYSTEMA • 2025 • 47 (31) Lemaitre R. INTRODUCTION Among the 88 genera currently in the family Paguridae Latreille, 1802, Catapagurus A. Milne-Edwards, 1880 is one of 56 (63.6%) of the genera in the family in which the males of the species are characterized by having sexual tubes that exhibit a remarkable variety of shapes and symmetry. Before Asakura’s (2001) revision of Catapagurus, only 11species of this genus were known from the world. Asakura’s study added eight new species which at the time he placed in the reinstated genus Hemipagurus Smith, 1881, and subsequently eight more have been described in various studies (McLaughlin 2002, 2004; Komai& Osawa 2009; Nucci& de Melo 2012; Komai& Rahayu 2021; Komai etal. 2022). An examination of selected samples of Catapagurus collected in the Western and Central South Pacific during French expedi - tions sponsored by the Muséum national d’Histoire naturelle, Paris (MNHN) and that were being studied by the late Patsy A. McLaughlin (see obituary in Lemaitre 2012), revealed the existence of two more new species that are herein described. With the description of these two new species, the genus now contains 30species and now ranks second among the genera with species having sexual tubes only to Catapaguroides A.MilneEdwards& Bouvier, 1892 which contains 32species. In addition, the studied samples contain specimens of C.franklinae McLaughlin, 2004 and C.tanimbarensis McLaughlin, 1997 from localities much eastward than previously known. MATERIAL AND METHODS The studied specimens remain deposited in the MNHN. Measurements of specimen size (+ 0.1mm) are of shield length, from the tip of rostrum to posterior midpoint of the shield. General terminology follows McLaughlin (2003), except for the use of “pleon” in lieu of “abdomen” in accord with Schram& Koenemann (2004), and thoracic sternites which are referred to only by the pereopodpairs between which they are ventrally positioned as per Felder etal. (2019). AbbreviAtions ovig ovigerous; stn station. Institutions MNHN Muséum national d’Histoire naturelle, Paris, USNM National Museum of Natural History, Smithsonian Institution, Washington D.C. SYSTEMATICS Family PAguridAe Latreille, 1802 Genus Catapagurus A.Milne-Edwards, 1880 Catapagurus poupini n.sp. (Figs1-3) urn:lsid:zoobank.org:act:76D5F6D5-9A08-4C40-A18D-0EC451E1938C tyPe mAteriAl. — Holotype. Marquesas Archipelago • ♂ 3.0mm; Hiva Oa island; MUSORSTOM 9; stn CP1212; 09°49.9’S, 139°02.2’W; 50-80m depth; shell sands with segment of Halimeda; 29.VIII.1997; MNHN-IU-2022-4536. Paratypes. Marquesas Archipelago • 2 ♂ 3.0-3.1mm; Hiva Oa Island; N.O. Alis, MUSORSTOM 9; stn CP1215; 49-62m depth; 09°49.5’S, 140°02.2’W; 29.VIII.1997; Bouchet, Dayrat, Richer de Forges legs; MNHN-IU-2022-4537 • 1 ♂ 2.6mm, 2ovig.♀ 2.5mm, 2.6mm; same data as for preceding; USNM 1441354 • 5 ♂ 2.1-2.7mm; Hiva Oa Island; N.O. Alis, MUSORSTOM9; stnCP1237; 95-305m depth; 09°42’S, 139°04’W; 31.VIII.1997; Bouchet, Dayrat, Richer de Forges legs; MNHN-IU-2022-4538 • 5 ♂ 2.1-2.8mm; same data as for preceding; USNM 1441353. other mAteriAl exAmined. — Marquesas Archipelago • 7 ♂ 1.9-3.0mm, 1 ♀ 3.0mm, 5ovig ♀ 2.4-2.8mm, [most dismembered]; Hiva Oa Island; N.O. Alis; MUSORSTOM 9; stnCP1212; 09°49.9’S, 139°02.2’W; 50-80m depth; 29.VIII.1997; Bouchet, Dayrat, Richer de Forges legs; USNM 1445847. etymology. — The specific epithet, poupini, is given to acknowledge the important contributions to our knowledge of tropical Pacific and Indian Ocean decapods by Dr Joseph Poupin (École navale, Brest, France). d istribution . — At present known only from the Marquesas Archipelago; 50-305m. hAbitAt. — Housing unrecorded, probably empty gastropod shells; on shell sands with segments of Halimeda algae. descriPtion of holotyPe Shield (Fig.1A) Broader than long; anterior margin between rostral lobe and lateral projections concave; anterolateral margins sloping; posterior margin roundly truncate; dorsal surface slightly convex, smooth, with few sparse tufts of moderate short setae. Rostral lobe broadly rounded, not reaching level of lateral projections. Lateral projections broadly triangular, terminating in sharp spine. Carapace lateral lobes narrow, reaching slightly beyond midlength of shield. Posterior carapace with poorly calcified median plate; cardiac sulci distinctly calcified, reaching posterior margin. Branchiostegites unarmed. Ocular peduncles (Fig.1A) Short, stout, 0.6-0.7 length of shield, broadened distally, corneal diameter 0.5-0.6 of peduncular length; ocular acicles, slender, terminally acute, reaching mid-length of ocular peduncles, mesial margins each with fringe of long setae, separated basally by twice basal width of one acicle. Antennular peduncles (Fig.1A) Overreaching distal corneal margins by 0.2-0.5 length of penultimate segments; dorsolateral surfaces and dorsolateral distal angles of ultimate segments each with few long setae; penultimate segment glabrous; basal segment with spine at ventromesial distal angle. Antennal peduncles (Fig.1A) Overreaching distal corneal margins by nearly entire length of ultimate (fifth) segments; fifth and fourth segments each with few scattered setae; third segment with ventrodistal angle produced, with blunt terminal spine; second segment with dorsolateral distal angle produced, terminating in strong spine,
753 New species and records of Catapagurus ZOOSYSTEMA • 2025 • 47 (31) dorsomesial distal angle with small spine; first segment with small spine on lateral face and at ventrodistal margin. Antennal acicles strong, reaching approximately mid-length of corneas, with few distal setae; mesial and lateral faces unarmed. Antennal flagella reaching to distal end of right chela; flagella naked. Mouthparts Not dissected. Third maxilliped with basis bearing four spinelike teeth; ischium crista dentata with six teeth (two spine-like proximally, four reduced and rounded distally), and prominent accessory tooth; merus with small dorsodistal spine. Chelipeds (Fig.2) Distinctly dissimilar, sparsely setose; left considerably slenderer than right; fingers bearing moderately long setae and denser than on rest of chelipeds, slightly crossed at tips, each terminating in small, blunt corneous claw. Right cheliped (Fig.2A, B) moderately stout, exceeding left cheliped in distal extension by more than length of dactyl of right. Dactyl approximately 0.6times as long as palm, with moderately long setae; dorsal surface convex, with scattered minute granules; dorsomesial margin not delimited, rounded and minutely granular; ventral and dorsal surfaces similar; A B C D A, B C, D D fig. 1. — Catapagurus poupini n. sp., holotype male 3.0 mm, Marquesas, Hiva Oa island, MUSORSTOM 9; stn CP1212, MNHN-IU-2022-4536: A, shield and cephalic appendages, dorsal; B, sternite, coxae of pereopods 1-5, and sexual tube, ventral; C, tip of sexual tube; D, uropods and telson, dorsal. Scale bars: A, B, 0.5 mm; C, D, 1.0 mm.
754 ZOOSYSTEMA • 2025 • 47 (31) Lemaitre R. cutting edge sinuous, armed with minute blunt teeth, distally forming a broadly rounded tooth, and sharp proximal tooth. Palm approximately as long as carpus; fixed finger with minutely granular surfaces (denser on lateral margin), cutting edge armed with minute blunt teeth, forming low, rounded tooth proximally; dorsal and ventral surfaces convex, dorsal surface with minute blunt to sharp granulation (stronger on dorsolateral and dorsomesial surfaces), dorsomesial and dorsolateral margins not delimited; ventral surface weakly granular. Carpus approximately equal to length of merus; dorsal, lateral and mesial surfaces distinctly granular, dorsolateral margin faintly delimited by small blunt spines proximally and small sharp spines distally, dorsomesial margin delimited by row of small spines increasing slightly in size distally; ventral surface AB CD fig. 2. — Catapagurus poupini n. sp., holotype male 3.0 mm, Marquesas, Hiva Oa island, MUSORSTOM 9; stn CP1212, MNHN-IU-2022-4536: A, right cheliped, dorsal; B, same, lateral; C, left cheliped, dorsal; D, same, lateral. Scale bar: 1.0 mm.
755 New species and records of Catapagurus ZOOSYSTEMA • 2025 • 47 (31) weakly granular. Merus triangular in cross-section; dorsal, lateral and mesial surfaces with granules or small spines stronger than on carpus, with distinct dorsodistal spine; ventrolateral and ventromesial margins well delimited by granules or small spines; ventral surface weakly granular. Ischium row of small spines on ventrolateral margin. Coxa unarmed. Left cheliped (Fig.2C, D). Distinctly shorter than right, reaching when fully extended approximately midlength of right palm, sparsely setose; fingers crossing at tips when closed, each terminating in sharp, inwardly curved corneous claw. Dactyl approximately 1.2 times as long as palm; dorsal, mesial, and ventral surfaces granular and with few scattered moderately long setae; cutting edge with row of small corneous teeth. Palm approximately 0.6 times as long as carpus; dorsal surface weakly convex; surfaces all bluntly to sharply granular, with scattered setae most abundant on fixed finger; fixed finger B A E F G A, D B, C, E, F G D C fig. 3. — Catapagurus poupini n. sp., holotype male 3.0 mm, Marquesas, Hiva Oa island, MUSORSTOM 9; stn CP1212, MNHN-IU-2022-4536: A, left pereopod 2, lateral; B, dactyl of same, lateral; C, same, mesial; D, left pereopod 3, lateral; E, dactyl of same, lateral; F, same, mesial; G, propodus and dactyl of left pereopod 4, lateral. Scale bars: A, D, 3.0 mm; B, C, E, F, 3.0 mm; G, 1.0 mm.
756 ZOOSYSTEMA • 2025 • 47 (31) Lemaitre R. lateral margin with few blunt or sharp small tubercles, cutting edge with small corneous teeth. Carpus slightly longer than merus, surfaces all minutely granular; dorsomesial margin weakly defined, rounded; dorsolateral margin delimited by row of small acute spines increasing somewhat in size distally. Merus triangular in cross-section, with row of few stiff bristles on dorsal margin; surfaces minutely granular; dorsodistal unarmed. Ischium with blunt dorsal spine, ventrolateral margin with row of few small spines. Coxa unarmed. Ambulatory legs (Fig.3) Pereopods2 and 3 similar except pereopod2 somewhat shorter. Dactyls weakly blade-shaped; in dorsal view slightly twisted inwardly, in lateral view somewhat arched ventrally; approximately 1.1 times as long as of propodi, each terminating in sharp corneous claw; dorsal margins with one row of short bristle-like setae, ventral margins unarmed; lateral faces with weak median longitudinal groove, lateral face of pereopod2 (Fig.3A-C) unarmed, lateral face of pereopod3 (Fig.3D-F) with one row of six short stout corneous spinules on ventral half; mesial faces weakly concave. Propodi with all surfaces, dorsal and ventral margins, minutely granular or with scattered setae. Carpi with all surfaces minutely granular, dorsodistal margins each with one row of few minute blunt spines. Meri with surfaces minutely granulated; dorsal margins each with moderately long setae proximally, armed distally with a row of distinct spines (rows of two spines on each meri on left side, and a row of 3-5spines on meri of right side). Pereopod4 (Fig.3G) With preungual process at base of claw; dactyl terminating in short, curved corneous claw; propodal rasp consisting of one row of ovate scales. Sternite of third pereopods(Fig.1B) with short anterior lobe having distal margin subdivided by shallow u-shaped sinus into two setose lobes. Male pereopod5, sexual tube, and pleopods(Fig.1B,C) With coxae (Fig.1B) roughly symmetrical; right coxa with sexual tube, left with gonopore obscured by fringe of setae; sexual tube (Fig.1B, C) long, slender, semitransparent, directed outward and curving over dorsal portion of pleon, terminally bilobed (one low lobe, and one slender, elongated lobe), spermatophores visible inside sexual tube; coxa of left with gonopore surrounded by dense fringe of setae. Male with three weakly developed left pleopods3-5, pleopods3 and 4 with rudimentary internal rami, pleopod5 uniramous. Females With pleopods 2-5 (pleopods 2-4 ovigerous). Uropods (Fig.1D) Asymmetrical; right protopod slightly produced posteriorly; exopods with moderately developed rasps. Telson (Fig.1D) Slightly asymmetrical, longer than broad, with distinct deep lateral indentations. Posterior lobes subtriangular, separated into two lobes by distinct, deep, U-shaped median cleft, outer margins broadly rounded with few short setae, apex of lobes with minute short bristles, inner margins unarmed. Color Unknown. remArks Among the 14 species of Catapagurus that have blade-shaped dactyls on pereopods2 and 3, this new species is most similar to three of those species in having weakly blade-shaped dactyls. Those three species are: C.fimbriatus Komai& Rahayu, 2021, C.insolitus Komai& Osawa, 2009, and C.tenuilamina Komai, Miller& Malay, 2022. However, that is where the similarity ends as there are a number albeit subtle characters that set C.poupini n.sp. apart from those other three. This new species differs from Catapagurus fimbriatus as follows: shape of shield (barely broader than long in the new species vs distinctly broader than long in C.fimbriatus); shape of cornea (roughly as broad as long in the new species vs distinctly broader than long in C.fimbriatus); teeth on cutting edge of dactyl of right chela (armed with minute blunt teeth proximally and a distal rounded tooth in the new species vs having two unequal prominent blunt teeth in C.fimbriatus); development of rasp on propodus of pereopod4 (consisting of a long row of small scales in the new species vs a short row of relatively large scales in C.fimbriatus); presence/absence of male gonopore on left coxa (present in the new species vs absent in C.fimbriatus); and, telson lateral indentations and shape of posterior lobes (with distinct lateral indentations, and U-shaped median cleft separating terminally blunt posterior lobes in the new species vs wide trapezoidal median cleft separating terminally acute posterior lobes in C.fimbriatus). This new species differs from Catapagurus insolitus as follows: the shape and armature of antennal acicles (slender and unarmed mesially in the new species vs stout and armed with distinct spines mesially in C.insolitus); length of pereopod4 preungual process (short and barely exceeding dactyl claw in the new species vs long and distinctly exceeding dactyl claw in C.insolitus); and, telson posterior lobes (separated by deep U-shaped median cleft in the new species vs separated by a a subrectangular median cleft in C.insolitus). This new species differs from Catapagurus tenuilamina as follows: length of pereopod4 preungual process (short and barely exceeding dactyl claw in the new species vs long and distinctly exceeding dactyl claw in C.tenuilamina); teeth on cutting edge of dactyl of right chela (armed with minute blunt teeth proximally and a distal rounded tooth in the new species vs one large blunt subtriangular tooth proximally and low round teeth distally in C.tenuilamana); armature of ventromesial margin of dactyl of pereopods2 and 3 (with one row of about six minute corneous spinules in the new species vs with one row of about 20minute spinules in C.tenuilamina); presence/absence of male gonopore on left coxa (present in the new species vs absent in C.tenuilamina); and terminal shape of sexual tube (bilobed with one low lobe slender, elongated vs bilobed with both lobes short and rounded in C.tenuilamina).
757 New species and records of Catapagurus ZOOSYSTEMA • 2025 • 47 (31) Catapagurus thysanodactylus n.sp. (Figs4-6) urn:lsid:zoobank.org:act:27CED4C9-8643-4F6E-BBFA-39007A1C2860 tyPe mAteriAl. — Holotype. Philippines • ♂ 3.0mm; Panglao Island, off San Isidro; PANGLAO 2004; stn T10; 09°33.4/33.8’N, 123°49.6/51.5’E; 117-124m depth; 15.VI.2004; mud and fine sand; 15.VI.2004; MNHN-IU-2022-4539. Paratypes. Philippines • ♂ 1.6mm; same station as holotype; MNHN-IU-2022-4545 • 7 ♂ 1.9-2.4mm, 6 ovig ♀ 1.8-2.3mm; between Panglao and Pamilacan islands; PANGLAO 2004; stn T27; 09°33.4’N, 123°51.0’E; 106-137m depth; fine sand and mud with echinoderms; 25.VI.2004; MNHN-IU-2022-4540. etymology. — The specific epithet is from the Greek thysanos, meaning fringe, in reference to the fringe of setae present on the dactyls of pereopods2 and 3. B A C D A B C, D fig. 4. — Catapagurus thysanodactylus n. sp., holotype male 3.0 mm, Philippines, Panglao Island, off San Isidro, PANGLAO 2004; stn T10, MNHN-IU-2022-4539: A, shield and cephalic appendages, dorsal; B, sternite, coxae of pereopods 1-5, and sexual tube, ventral; C, tip of sexual tube; D, uropods and telson, dorsal. Scale bars: A, 1.0 mm; B-D, 0.5 mm.
764 ZOOSYSTEMA • 2025 • 47 (31) Lemaitre R. remArks This species was previously known only from the Kai and Tanimbar Islands, Indonesia (McLaughlin 1997). The current record extends the range of this species approximately 3200km eastwards to the Solomon Islands. Acknowledgements The material of Catapagurus used here was being worked on by DrPatsy A. McLaughlin shortly before her death on April of 2011. The specimens were found by the author in her personal laboratory and were part of a major review of Catapagurus which she regrettably never lived to finish. She had recognized several new species while working on the material, much of it collected during French expeditions led by the MNHN (VOLSMAR ( https://doi.org/10.17600/89005011); SALOMON 2 (https://doi.org/10.17600/4100090); BOA 1 (https://doi.org/10.17600/5100060); BORDAU 1 (https:// doi.org/10.17600/99100020); MUSORSTOM 9 (https://doi. org/10.17600/97100020) to the Western Pacific, all of which has been returned to that Museum. The PANGLAO Marine Biodiversity Project including the cruise PANGLAO 2024, was a joint project of MNHN, Paris (PI: Philippe Bouchet) and University of San Carlos, Cebu City (PI: Danilo Largo), funded by grants from the Total Foundation, the French Ministry of Foreign Affairs, and the Asean Regional Center for Biodiversity Conservation (ARCBC), and operating under a permit from the Philippine Bureau of Fisheries and Aquatic Resources (BFAR). Unfortunately, it was difficult to reconcile the numerous specimens and labels with her manuscript notes and observations left scattered in her laboratory, except for the specimens of the two new species described herein. It seemed appropriate to maintain the specific epithets she had chosen for these two new species, even though she never wrote descriptions or prepared illustrations. The author did not want to leave the work unfinished, and thanks her posthumously. 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Annals and Magazine of Natural History(5)7: 143-146. https://doi.org/10.1080/00222938109459488 Submitted on20 November 2024; accepted on 30 April 2025; published on 3 December 2025.