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Latin american journal of aquatic research

On-line version ISSN 0718-560X

Lat. Am. J. Aquat. Res. vol.45 no.2 Valparaíso May 2017 

Research Article


Taxonomy of Pinnotheres bipunctatus Nicolet, 1849 with a distributional checklist of the Pinnotheridae of Chile and Peru, and a list of the Crustacea described by Hercule Nicolet in the Atlas of the Physical and Political History of Chile


Ernesto Campos1

1 Facultad de Ciencias, Universidad Autonoma de Baja California, Baja California, Mexico

Corresponding author: Ernesto Campos (
Corresponding editor: Ingo Wehrtmann

ABSTRACT. Because the male holotype of Pinnotheres bipunctatus possesses a carpus of the third maxilliped larger than the propodus, the dactylus disto-medially inserted on the ventral margin of this latter article extending far beyond its tip, and a laterally expanded telson, wider than the sixth abdominal somite, it is excluded from Pinnotheres and assigned to Pinnaxodes (type species P. chilensis). Although males of these species can morphologically be separated, the holotype of P. bipunctatus resembles a juvenile, consequently, a categorical taxonomic distinction between this species and P. chilensis will require a comparative morphological study of the preadult stages of the latter species. A distributional checklist and host of the Pinnotheridae recorded for the Peru-Chile region is included, along with a list and the taxonomic status of the five genera and 53 species of Crustacea described by Hercule Nicolet in the "Physical and Political History of Chile".

Keywords: Pinnotheridae, Pinnaxodes, Echinodermata, Claudio Gay, Chile.



Ongoing taxonomic studies on the pinnotherid crabs of the Pacific coast of South America have prompted a reassessment of the taxonomy of the species Pinnotheres bipuncatus Nicolet, 1849. This small crab was the only species of Pinnotheridae described by the Swiss naturalist Hercule Nicolet in the chapter of Crustacea (Zoology, section 3) of the "Physical and Political History of Chile" by Claudio Gay (1849). The morphology of P. bipunctatus was analyzed and compared with the type species of Pinnotheres Bosc, 1802, P. pisum (Linnaeus, 1767), as well as other genera of Pinnotheridae of the Eastern Pacific (Campos, 2009, 2016). The long dactylus of the third maxilliped that overreaches the tip of the propodus and the subtriangular telson wider than the sixth abdominal somite are unique features that clearly support that P. bipunctatus does not belong to Pinnotheres. However, it closely resembles Pinnaxodes chilensis (H. Milne-Edwards, 1837), type species of the genus Pinnaxodes Heller, 1865. The shared morphology between P. bipunctatus and P. chilensis supports the conclusion that both are congenerics, but also raises the hypothesis that both species probably were described on the basis of two different developmental stages (juvenile and adult, respectively) of the same species.

A distributional checklist of the Pinnotheridae recorded for the Peru-Chile region is presented along with a taxonomically updated list of the species of Crustacea described by H. Nicolet.


The checklist was prepared using available literature and specialized databases (Table 1). Because the holotype of P. bipunctatus is not extant, I relied on the original description and figures published by Nicolet (1849). The description originally written in Spanish was translated to English by Rathbun (1918). Additionally, voucher specimens of Pinnaxodes chilensis and Pinnotheres pisum were studied from material deposited in the Laboratorio de Invertebrados of the Facultad de Ciencias, Universidad Autónoma de Baja California (UABC) and the Smithsonian Institution (USNM). Drawings were made with the aid of a camera lucida attached to a stereoscopic microscope. Editing of the drawings was performed using the Adobe Illustrator CS and Adobe Photoshop CS computer programs.


Table 1. Distributional checklist and hosts of the Pinnotheridae of Peru and Chile.



Systematics Account

Pinnotheridae De Haan, 1833

Pinnaxodes Heller, 1865

Pinnaxodes chilensis (H. Milne-Edwards, 1837)

Restricted synonymy: see Schmitt et al. (1973) for a complete synonymy.

Pinnotheres chilensis H. Milne Edwards, 1837: 33, (Atlas): pl. 10, fig. 2; H. Milne Edwards & Lucas, 18421844: 23; Nicolet in Gay, 1849: 155; Rathbun, 1918: 175-177, pl. 38, figs. 1-7; Garth, 1957: 70, 85-88, 92, fig. 9A-F; Schmitt et al, 1973: 33-34; Takeda & Masahito, 2000: 99-112, fig. 1A, 2C-D; Ng & Manning, 2003: 914-916, Figs. 7a, 7b.

Fabia chilensis Dana, 1852: 383 (type locality, near Valparaíso, Chile).

Pinnaxodes hirtipes Heller, 1865: 68, pl. 6, Fig. 2 (type locality, Ecuador)


Carapace soft and yielding in female, firm, parchmentlike in male. Palm of female elongate. Front slightly produced, deflexed, divided by a shallow medial sulcus. Outer maxillipeds placed nearly longitudinally; merus and ischium fused, a demarcation line sometimes visible between them; palpus three segmented, carpus slightly longer than subtrapezoidal propodus, dactylus articulated disto-medially on ventral margin of subtrapezoidal propodus, overreaching it considerably. Ambulatory legs 1-4 (pereiopods 2-5) similar in females, somewhat unequal and longer in males, dactyli slender in both sexes, longer in males, third longest in male, fourth longest in female; third ambulatory leg longest in males, second and third ambulatory legs subequal and longest in females. Abdomen of six somites and telson free, that of female wide, long, that of male narrow at base, tapering from third to sixth somite which has lateral margins concaves, telson laterally expanded, wider than somite six.

Type species and host

By original designation and monotypy, Pinnaxodes hirtipes Heller, 1865 (= Pinnotheres chilensis H. Milne-Edwards, 1837), gender masculine. Associated with Echinodermata, Echinoidea: Echinometridae, Caenocentrotus gibbosus (L. Agassiz, in L. Agassiz & Desor, 1846); Parechinidae, Loxechinus albus (Molina, 1782); Arbaciidae, Tetrapygus niger (Molina, 1782) (Fenucci, 1967; Schmitt et al., 1973).

Distribution of type species

Ecuador (type locality) to Port Otway, Chile; Chiloé Island; Tierra del Fuego; Galapagos Islands (Fenucci, 1967; Schmitt et al., 1973). To a depth of one fathom (1.83 m) (Rathbun, 1918); recent commercial catches of L. albus infested with P. chilensis came from a medial depth of 25 m (range 0-40 m) (Runil-Ojeda, 2014). Specimens collected during the Albatross expedition (see below) were caught in 104 m depth.

Other species included in Pinnaxodes Heller

The following species have been included in this genus: P. floridensis Wells & Wells, 1961 (off North Carolina to Georgia; northwest Florida: Williams, 1984); P. gigas Green, 1992 (Gulf of California and West coast of Baja California: Campos et al., 1998, Campos, 2016); P. major Ortmann, 1894 (China, South Korea, Japan, and the Russian Far East: Marin, 2014); P. mutuensis Sakai, 1939 (northern Japan: Takeda & Masahito, 2000) and P. tomentosus Ortmann, 1894 (Brazil: Ortmann, 1894; Melo & Boehs, 2004).

Pinnaxodes bipunctatus (Nicolet, 1849) new combination


Pinnotheres bipunctatum Nicolet in Gay, 1849: 155156 (type locality, San Carlos de Chiloe [= Ancud]); 1854, Atlas, Crustacea, pl. 1, fig. 2a-c; Milne-Edwards, 1853: 219; Rathbun, 1910: 587; Tesch, 1918: 286 (listed); Silas & Alagarswami, 1967: 1196, 1222 (listed).

Pinnotheres bipunctatus, Philippi, 1894, 372 (listed); Porter, 1909: 249 (in footnote); 1911: 446 (in footnote); Rathbun, 1918: 78, 159; Garth, 1957: 70, 92; Schmitt et al, 1973: 40; Rodríguez, 1993: 47.

Description (slightly modified from Nicolet, 1849 and Rathbun, 1918)

Carapace wider than long, curved along sides, narrowing slightly toward back, posterior border straight, two rather large punctate in middle. Front quadrilateral, transverse, prominent beyond curve of anterolateral borders, its anterior margin broad and slightly hollowed out, its middle occupied by a longitudinal depression bordered on each side by raised, rounded, and forward-pointing projection. Orbits small, but deep. Chelipeds and ambulatory legs robust, compressed; chelipeds shorter than pereiopods 2-4; hand short, wide, and nearly quadrate, dactylus wider than pollex and strongly curved. Ambulatory legs (pereiopods 2-5) covered with very short, coarse hair, scarcely visible; dactyli strong, curved, and with a sharp claw. Abdomen narrow, elongate; telson sub-triangular, wider than somite six. Posterior part of body rough, covered with spine-like hairs, some small ones on the inner margin of outer maxillipeds.

Distribution and host

Known only from the type locality, San Carlos de Chiloe (Ancud), Chile. Probably in sea urchins (Nicolet, 1849).

Material examined

None. The male holotype is not extant.


Shiny flavo (yellow-gold) (Nicolet, 1849).


Length of male carapace, 1 to 2 lines. A line is an obsolete French unit of length equal to 2.3 mm (Cardarelli, 2004). According to figure 2a of Nicolet (1854), carapace length = 3.6 mm, carapace wide = 4.2 mm.

Additional material examined

Pinnotheres pisum, "coast of France" UABC; Pinnaxodes chilensis, Albatross R/V, station 2786 (46°46'00"S, 75°16'30"W), 104 m depth, Peninsula Taitao, Gulf of Penas, 8 Feb 1888, 1 male, USNM 22112; same, 1 female, USNM 49238; 2 females, "coast of Chile" in Loxechinus albus (Molina, 1782), UABC.


Garth (1957) analyzed the taxonomic status of Pinnotheres bipunctatus and pointed out the male described by Nicolet (1849) may belong to P. politus Smith, 1870 (now in Calyptraeotheres Campos, 1990). Although both species have a similar habitus (Figs. 1a-1b), remarkable differences exist in the third maxilliped and abdomen that allow both rejecting this idea and the placement of this species in Calyptraeotheres.


Figure 1. Male, dorsal view: a) Calyptraeotheres politus
(Smith, 1870), Seno Reloncavi, Punta Pilluco (Pelluhuin),
Chile; carapace width: 3.7 mm; carapace length: 3.6 mm,
b) Pinnaxodes bipunctatus (Nicolet, 1849) San Carlos de
Chiloe, Chile; carapace width: 4.16 mm; carapace length:
3.6 mm (Nicolet, 1854). c) Pinnaxodes chilensis (H. Milne
Edwards, 1837) Port San Pedro, Chiloe Island, Chile;
carapace width: 7.3 mm; carapace length: 6.6 mm (Garth,

1957). a, c) from Garth, (1957); b) from Nicolet (1854).


The limpet crab C. politus has a third maxilliped with a minute dactylus inserted subdistally on the ventral margin of propodus (Figs. 2a, 2c) and a subcircular telson (Fig. 2b). Contrarily, P. bipunctatus has a long dactylus inserted disto-medially on ventral margin of propodus and overreaching the tip of this latter article, and a subtriangular telson, laterally expanded, which is wider than the sixth somite (Figs. 3a-3b).


Figure 2. Calyptraeotheres politus (Smith, 1870), a) third
maxilliped, b) abdomen, Seno de Reloncavi, Punta Pilluco
(Pelluhuin), Chile, carapace width: 3.7 mm, carapace length:
3.6 mm, not a scale; c) third maxilliped, scale: 0.78 mm.
(Linnaeus, 1767): d, f) outer view third
maxilliped, d) coast of France (UABC), carapace width:
8.2 mm, carapace length: 7.3 mm, not a scale, f) off
Senigallia, Italy (USNM 205776) not a scale; e) abdomen,
coast of France (UABC), length: 3.4 mm. a-b) from Garth
(1957); c) from Campos (1999); f) from Manning (1993a).


Figure 3. Pinnaxodes bipunctatus (Nicolet, 1849), new combination,
San Carlos de Chiloé, Chile, carapace wide: 4.16 mm; carapace length:
3.6 mm (Nicolet, 1854): a), third maxilliped; b) abdomen. Pinnaxodes
(H. Milne Edwards, 1837): c) abdomen; d-f) third maxilliped:
c-d) Port San Pedro, Chiloé Island, Chile; e) Port Otway, Patagonia;
f) Chile. a,b, e) not a scale; c) X21; D, X57.6; f) scale: 1 mm. a-b
from Nicolet (1854); c-d) from Garth (1957); e) from Rathbun (1918);
f) from Ng & Manning (2003).


In addition, a detailed analysis of the description and figures of P. bipunctatus reveals that this species clearly does not belong to the genus Pinnotheres. The principal differences are observed in the previously described third maxilliped and abdomen. Members of the genus Pinnotheres have a carpus shorter than the propodus and a digitiform dactylus that is proximally inserted on the ventral margin of the propodus, falling short of the tip of this latter article (Figs. 2d, 2f), while the abdomen has a subcircular telson and is neither laterally expanded nor wider than the sixth somite (Fig. 2e). The morphological features and the original statement that P. bipunctatus probably was collected in a sea urchin support its exclusion from Pinnotheres, a symbiont of bivalves (Manning, 1993b). Nevertheless, these features allow the conclusion that P. bipunctatus is a member of the genus Pinnaxodes Heller. Both species, P. bipunctatus Nicolet new combination and P. chilensis (type species of Pinnaxodes), share a similar third maxilliped with a long dactylus that overreach the tip of the propodus (Figs. 3a, 3d, 3e, 3f) and a subtriangular telson wider than the sixth abdominal somite (Figs. 3b, 3c). Additional shared features include a produced and emarginate front, a carapace with two large punctate and a medial sulcus that arises in the front and extends to the gastric region, and the ambulatory legs 1-3 (pereiopods 2-4) are longer than the chelipeds. Moreover, the host of P. bipunctatus is presumably a sea urchin; if so, it would be a similar host as that of P. chilensis.

The main morphological differences between the male of P. bipunctatus and P. chilensis include the ambulatory legs and their dactyli, proportionally shorter in the former species and notoriously larger in P. chilensis (Figs. 1b-1c). However, the male described by Nicolet is a very small specimen with only 4.2 mm carapace width, while males described for P. chilensis are about twice in size (Rathbun, 1918; Garth, 1957). This may suggest that the above-mentioned differences might be related to the size. The shared features and the morphological differences presumably associated to the size allow to hypothesize that the male described by Nicolet is an immature stage of P. chilensis; however, a categorical conclusion can currently not established and is awaiting a morphological study on the postlarval development of this species.

With the inclusion of P. bipunctatus in Pinnaxodes, a total of 10 species in five genera of Pinnotheridae are known for the temperate coasts of Peru and Chile (Peruvian-Chilean Province sensu Retamal & Moyano, 2010) (Table 1). Most of these species were described between 1836 and 1907 (seven species) and only three new species of Pinnixa White, 1846 were added by Rathbun (1935) and Garth (1957). The genus Pinnixa now represents the most diverse taxa of this family along the Pacific temperate coast of South America (Schmitt et al., 1973). In addition, Melzer & Schwabe (2008) recorded three juveniles of a pinnotherid harbored in the chiton Tonicia chilensis (Frembly, 1827) collected in Muelle Dichato, Chile (36°33'S, 72° 56'W, 1 m). According to these authors, the third maxilliped morphology of these juveniles suggests a closer relationship with the genus Orthotheres Sakai, 1969. However, the morphology of the third maxilliped of these minute crabs supports a different generic assignment that will be discussed elsewhere.

Species described by Hercule Nicolet

Claudio Gay (1847) in the preface of the "Physical and Political History of Chile" (Zoology, section 1) explicitly pointed out that the entomologist Hercule Nicolet was in charge of the section of Crustacea (Zoology, section 3) of his monumental treatise. This statement fully complies with article 50.1 and 50.1.1 of the International Code of Zoological Nomenclature that deals with the identity of authors. Nicolet (1849) described five genera and 53 species new to the fauna of Chile (Table 2) of which only the genus Orchestoidea Nicolet, 1849 and nine species remained without taxonomic changes. These include one Decapoda (Pagurus villosus Nicolet, 1849), two Amphipoda (Orchestia gayi Nicolet, 1849, Orchestoidea tuberculata Nicolet, 1849), four Isopoda (Oniscus armatus Nicolet, 1849, Porcellio liliputanus Nicolet, 1849, Sphaeroma propinqua Nicolet, 1849, S. gayi Nicolet, 1849) and two Tanaidacea (Tanais macrocheles Nicolet, 1849, T. gayi Nicolet, 1849). Additionally, 12 species remained valid but have been transferred to another genus, while the remaining 32 species included junior synonyms or species poorly known taxonomically and have been considered incertae sedis, nomen dubium or species inquirenda. The second decapod species and the only pinnotherid described by Nicolet is Pinnaxodes bipunctatus new combination, which is herein considered a valid species; however, it remains unclear whether it is a juvenile and consequently a junior synonym of P. chilensis as previously discussed.


Table 2. Genera and species of Crustacea described by Hercule Nicolet in Claudio Gay
(1849). Original taxonomic name at left is followed by the currently name accepted.
Sources of taxonomic information are: Martens & Behen (1994); Leistikow & Wäegele
(1999); Guerra-Garcia & Thiel, (2001); Schmalfuss (2003); Gonzalez et al. (2008);
Hayes et al. (2012); Kotov et al. (2013); Brandão et al. (2015); Mees et al. (2015);
Walter & Boxshall (2015); Anonymous (2016); Boxshall et al. (2016).



I remain indebted with Ingo Wehrtmann, Escuela de Biología, Universidad de Costa Rica and Hans Bertsch (Instituto de Investigaciones Oceanológicas, UABC) for the revision of an early draft of this manuscript, and to Alma Rosa de Campos for preparing the figures. My enduring and deep gratitude to the late Raymond B. Manning for having lent and donated material for this study. The Mexican Network for study of Exotic Species (SEP-PRODEP) has supported this research.



Agassiz, L. 1841. Monographies d'Échinodermes vivans et fossiles. Échinites. Famille des Clypéasteroides. 2 (Seconde Monographie). Des Scutelles, Neuchâtel, Switzerland, i-iv, 1-151: pp. 1-27.         [ Links ]

Agassiz, L. 1846. Caenocentrotus gibbosus. In: L. Agassiz & P.J.E. Desor (eds.). Catalogue raisonné des familles, des genres, et des espèces de la classe des échinodermes. Ann. Sci. Nat. Troisième Série, Zool., 6: 305-374.         [ Links ]

Anonymous 2017. Global Name Index. []. Reviewed: 9 January 2016.         [ Links ]

Born I. Von. 1778. Index rerum naturalium Musei Cœsarei Vindobonensis. Pars Testacea. Verzeichnis der natürlichen Seltenheiten des k.k. Naturalien Cabinets zu Wien. Erster Theil. Schalthiere. [1-40], 1-458, [182], Vindobonœ, (Kraus).

Bosc, L.A.G. 1802. Histoire naturelle des Crustacés, contenant leur description et leurs moeurs, avec figures dessinées d'après nature. Deterville, Paris, 1: 258 pp.         [ Links ]

Bott, R.1955. Dekapoden aus EI Salvador. Senck. Biol., 36: 45-72.         [ Links ]

Boxshall, G.A., J. Mees, M.J. Costello, F. Hernandez, N. Bailly, N. Boury-Esnault, S. Gofas, T. Horton, (et al.). 2016. World Register of Marine Species. []. Reviewed: 9 January 2017.         [ Links ]

Brandão, S.N., M.V. Angel, I. Karanovic, A. Parker, V. Perrier, B. Sames & M. Yasuhara. 2015. World Ostracoda Database. []. Reviewed: 9 January 2016.         [ Links ]

Campos, E. 1990. Calyptraeotheres, a new genus of Pinnotheridae for the limpet crab Fabia granti Glassell, 1933 (Crustacea, Brachyura). Proc. Biol. Soc. Wash., 103: 364-371.         [ Links ]

Campos, E. 1999. Inclusion of the austral species Pinnotheres politus (Smith, 1869) and Pinnotheres garthi Fenucci, 1975 within the genus Calyptraeotheres Campos, 1990 (Crustacea: Brachyura: Pinnotheridae). Proc. Biol. Soc. Wash., 112: 536-540.         [ Links ]

Campos, E. 2009. A new species and two new genera of pinnotherid crabs from the northeastern Pacific Ocean, with a reappraisal of the subfamily Pinnotherinae de Haan, 1833 (Crustacea: Brachyura: Pinnotheridae). Zootaxa, 2022: 29-44.         [ Links ]

Campos, E. 2016. The Pinnotheridae of the northeastern Pacific (Alaska to Mexico): zoogeographical remarks and new bivalve hosts (Crustacea, Brachyura, Pinnotheridae). Zootaxa, 4170(2): 311-329.         [ Links ]

Campos, E., A.R. de Campos & J.A. de León-González. 2009. Diversity and ecological remarks of ectocom-mensals and ectoparasites (Annelida, Crustacea, Mollusca) of echinoids (Echinoidea: Mellitidae) in the Sea of Cortez, Mexico. Parasitol. Res., 105: 479-487.         [ Links ]

Campos, E., V. Díaz & J.A. Gamboa-Contreras. 1998. Notes on distribution and taxonomy of five poorly known species of pinnotherid crabs from the eastern Pacific (Crustacea: Brachyura: Pinnotheridae). Proc. Biol. Soc. Wash., 111: 372-381.         [ Links ]

Cardarelli, F. 2004. Encyclopaedia of Scientific units, weights and measures: their SI equivalences and origins. Springer, London, 848 pp.         [ Links ]

Coppard, S.E. 2016. A new genus of mellitid sand dollar (Echinoidea: Mellitidae) from the eastern Pacific coast of the Americas. Zootaxa., 4111(2): 158-166.         [ Links ]

Dana, J.D. 1852. Crustacea. Part I. In: United States Exploring Expedition. During the years 1838, 1839, 1840, 1841, 1842. Under the command of Charles Wilkes, U.S.N. C. Sherman, Philadelphia, 13: 685 pp.         [ Links ]

De Haan, W. 1833-1850. Crustacea, plates A-J, L-Q 1-55, circ. tab. 2 In: Fauna Japonica sive Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent. Suscepto, Annis 1823-1830 Collegit, Notis, Observationibus et Adumbrationibus Ilustravit. i-xvii, i-xxxi, ixxvi (P.F. v. Siebold, ed.), Lugduni-Batavorum, Leiden, 243 pp.         [ Links ]

Fabricius, J.C. 1775. Systema Entomologiae, sistens Insectorum Classes, Ordines, Genera, Species, adjectis Sysnonymis, Locis, Descriptionibus, Observationibus. Kortii, Flensburgi et Lipsiae, 832 pp.         [ Links ]

Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America to the Galapagos,to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commission steamer "Albatross", during 1891, Lieut. Commander Z.L. Tanner, U.S.N., commanding. VI. Preliminary descriptions of new species of Crustacea. Bull. Mus. Comp. Zool. Harvard, 24: 149-220.         [ Links ]

Fenucci, J.L. 1967. Contribución al conocimiento del crustáceo decápodo braquiuro Pinnaxodes chilensis M. Edwards, comensal de Loxechinus albus (Molina) (Echinodermata, Echinoidea). Physis, Buenos Aires, 27: 125-133.         [ Links ]

Fischer, S. 1860. Beiträge zur Kenntniss der Entomos-traceen. Aus den Abhandlungen der k. bayer. Akademie d.W.II.Cl.VIII.Bd.III.Abth., 8, Abt. 3: 645-682.         [ Links ]

Frembly, J. 1827. A description of several new species of chitones, found on the coast of Chili, in 1825, with a few remarks on the method of taking and preserving them. Zool. J., 3: 193-205.         [ Links ]

Gallardo, C.S. 1979. Especies gemelas del género Crepidula (Gastropoda : Calyptraeidae) en la costa de Chile: una redescripción de Crepidula dilatata Lamarck y descripción de Crepidula fecunda n. sp. Stud. Neotrop. Fauna Environ., 14: 215-226.         [ Links ]

Garth, J.S. 1957. Reports of the Lund University Chile Expedition 1948-49 29. The Crustacea Decapoda Brachyura of Chile. Lunds Univers. Årsskrift, N.F. Avd, 2., 53: 1-129.         [ Links ]

Gay, C. 1847. Historia Física y Política de Chile, Zoología 1, Paris and Santiago, 496 pp.         [ Links ]

Gay, C. 1849. Historia Física y Política de Chile, Zoología 2, Paris and Santiago, 547 pp.         [ Links ]

González, E.R., P.A. Haye, M.-J. Balanda & M. Thiel 2008. Lista sistemática de especies de peracáridos de Chile (Crustacea, Eumalacostraca). Gayana, 72(2): 157-177.         [ Links ]

Green, T.M. 1992. Pinnaxodes gigas, a new species of pinnotherid crab from the Gulf of California (Decapoda: Brachyura: Pinnotheridae). Proc. Biol. Soc. Wash., 105: 775-779.         [ Links ]

Guérin-Méneville, F.E. 1835. Observations sur les Porcellanes. (F.-E.Guérin). Bull. Soc. Sci. Nat. France, pp. 115-116.         [ Links ]

Guérin-Méneville F.E. 1843. Crustacés. In: G. Cuvier (ed.). Iconographie du Regne animal de G. Cuvier, ou representation d'apres nature de l'une des especes les plus remarquable et souvent non encore figurees de chaque genre d'animaux. Volume 3, Paris, Bailliere, pp. 1-48.         [ Links ]

Guerra-García, J.M. & M. Thiel. 2001. The caprellid fauna (Crustacea: Amphipoda: Caprellidea) from the coast of Coquimbo, northern-central Chile, with a taxonomic key for species identification. Rev. Chil. Hist. Nat., 74: 873-883.         [ Links ]

Hayes, P., J.L. Justine & G.A. Boxshall. 2012. The genus Caligus Müller, 1785 (Copepoda: Siphonosto-matoida): two new species from reef associated fishes in New Caledonia, and some nomenclatural problems resolved. Zootaxa, 3534: 21-39.         [ Links ]

Heller, C. 1865. Crustaceen. In: Reise der österreichischen Fregatte "Novara" um die Erde in den Jahren 18571859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zool. Theil, Zweiter Band III. Abteilung, pp. 1-280.         [ Links ]

Hendrickx, M.E. 1995. Checklist of brachyuran crabs (Crustacea: Decapoda) from the Eastern Tropical Pacific. Bull. Inst. Roy. Sci. Nat. Bruxelles, 65: 125-150.         [ Links ]

Kingsley, J.S. 1878a. A synopsis of the North American species of the genus Alpheus. Bull. U.S. Geol. Geogr. Survey Terr., 4: 189-199.         [ Links ]

Kotov, A., L. Forró, N.M. Korovchinsky & A. Petrusek. 2013. FADA Cladocera: world checklist of freshwater Cladocera species (version Jan. 2013). In: Y. Roskov, L. Abucay, T. Orrell, D. Nicolson, T. Kunze, C. Flann, N. Bailly, P. Kirk, T. Bourgoin, R.E. DeWalt, W. Decock & A. De Wever (eds.). Species 2000 & ITIS Catalogue of Life, [] Reviewed: 15 January 2016.         [ Links ]

Kroyer H. 1845. Karcinologiske Bidrag. Naturh.Tidsskr., 1, 283-345, 403, 453-638 + pls 2-3, 6-7.         [ Links ]

Lamarck, J.B. 1822 Histoire naturelle des animaux sans vertèbres. Published by the author, Paris, 7: 711 pp.         [ Links ]

Leach, W.E. 1814. Crustaceology. In: D. Brewster (ed.). The Edinburgh encyclopaedia 7. Blackwood, Edinburgh, pp. 383-437.         [ Links ]

Leistikow, A. & J.W. Wäegele. 1999. Checklist of the terrestrial isopods of the new world (Crustacea, Isopoda, Oniscidea). Rev. Bras. Zool., 16: 1-72.         [ Links ]

Linnaeus, C. 1767. Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Editio duodecima, reformata. Regnum Animale. Tom. I. Pars II. Laurentii Salvii, Holmiae, 1327 pp.         [ Links ]

Manning, R.B. 1993a. West African pinnotherid crabs subfamily Pinnotherinae (Crustacea, Decapoda, Brachyura). Bull. Mus. Natl. Hist. Nat., Paris, (4)15A (1-4): 125-177.         [ Links ]

Manning, R.B. 1993b. Three genera removed from the synonymy of Pinnotheres Bosc, 1802 (Brachyura: Pinnotheridae). Proc. Biol. Soc. Wash., 106: 523-531.         [ Links ]

Martens, K. & F. Behen. 1994. A checklist of the recent non-marine ostracods (Crustacea, Ostracoda) from the inland waters of South America and adjacent islands. Trav. Scien. Musée Nat. Hist. Nat., Luxembourg, 22: 1-81.         [ Links ]

Marin, I.N. 2014. Finding of the pea crab Pinnaxodes mutuensis Sakai, 1939 (Crustacea: Decapoda: Pinnotheridae) in an unusual host in Busse Lagoon, southern Sakhalin. Russ. J. Mar. Biol., 40: 486-489.         [ Links ]

Mees, J., G.A. Boxshall, M.J. Costello, F. Hernandez, S. Gofas, B.W. Hoeksema et al. 2015. World Register of Marine Species (WoRMS). WoRMS Editorial Board. []. Reviewed: 15 January 2016.         [ Links ]

Melo, G.A.S. & G. Boehs. 2004. Rediscovery of Holothuriophilus tomentosus (Ortmann) comb. nov. (Crustacea, Brachyura, Pinnotheridae) in the Brazilian coast. Rev. Braz. Zool., 21: 229-232.         [ Links ]

Melzer, R.R. & E. Schwabe. 2008. First record of an early pea crab stage on Tonicia chilensis (Frembly, 1827) (Mollusca, Polyplacophora). Zootaxa, 1919: 58-60.         [ Links ]

Michelin, H. 1858. Revue des espèces connues et nouvelles du genre Mellita, famille des clypeastroides. Rev. Mag. Zool., Ser. 2, 10: 358-364.         [ Links ]

Miers, E.J. 1881. Revision of the Idoteidae, a family of sessile-eyed Crustacea. J. Linn. Soc. London, 16: 1-88.         [ Links ]

Milne-Edwards, H. 1837. Histoire naturelle des Crustacés, comprenant l'anatomie, la physiologie et la classification de ces animaux, 2:532 pages. Atlas [1834, 1837, 1840]: 32 pages, plates 1-14, 14bis, 15-25, 25bis, 2642 Paris.         [ Links ]

Milne-Edwards, H. 1853. Observations sur les affinites zoologiques et la classification naturelle des Crustacés. Ann. Sci. Nat., 3è Sér., 20: 163-228.         [ Links ]

Milne-Edwards, H. & H. Lucas. 1842-1844. Crustacés. In: A. d'Orbigny (ed.). Voyage dans l'Amerique méridionale (le Brésil, la république orientale de l'Uruguay, la république Argentine, la Patagonie, la république du Chili, la république de Bolivia, la république du Pérou), exécutée pendant les années 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833 P. Bertrand, Ve Levrault, Paris, Strasbourg, pp. 1-37.         [ Links ]

Molina, G.I. 1782. Saggito sulla storia naturale del Chili. 4, Animali del Chili, Bolonga, 367 pp.         [ Links ]

Moscoso, V. 2012. Catálogo de crustáceos decápodos y estomatópodos del Perú. Bol. Inst. Mar Perú, 27: 1-209.         [ Links ]

Müller, M. 1852. Observationes anatomicae de vermibus quibusdam maritimis. Diss. Berlin, pp. 14-22.         [ Links ]

Nobili, G. 1901. Decapodi raccolti dal Dr. Filippo Silvestri nell'America meridionale. Boll. Mus. Zool. Anat. Comp. Torino, 16(402): 1-16.         [ Links ]

Ng, P.K.L. & R.B. Manning. 2003. On two new genera of pea crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura: Pinnotheridae) from the IndoWest Pacific, with notes on allied genera. Proc. Biol. Soc. Wash., 116: 901-919.         [ Links ]

Nicolet, H. 1849. Crustacea. In: C. Gay (ed.). Historia física y política de Chile, Zoología, tomo 3, Paris and Santiago, pp. 115-318.         [ Links ]

Nicolet, H. 1854. Crustáceos (Plate 1-4). In: C. Gay (ed.). Atlas de historia física y política de Chile, Zoología, Tomo 2, Paris and Santiago, unnumbered pages.         [ Links ]

Ortmann, A.E. 1894. Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen. VIII Theil. Abtheilung: Brachyura (Brachyura Genuina Boas). III. Unterabtheilung: Cancroidea, 2. Section: Cancrinea, 2. Gruppe: Catametopa. Zool. Jahrb. Allg. Zool. Syst., Geogr. Biol. Thiere, 7: 683-772.         [ Links ]

Paredes, C. & F. Cardoso. 2007. La Familia Calyptraeidae en el Perú (Gastropoda: Caenogastropoda). Rev. Peru. Biol., 13(3): 177-184.         [ Links ]

Philippi, R.A. 1894. Dos palabras sobre la sinonimia de los crustáceos, decápodos, braquiuros o jaivas de Chile. An. Univ. Chile, 87: 369-376.         [ Links ]

Poeppig, E. 1836. Crustacea Chilensia nova aut minus nota descripsit. Arch. Naturgesch., 2(1):133-145.         [ Links ]

Porter, C.E. 1909. Materiales para la fauna carcinológica de Chile VII. La familia Pinnotheridae. Rev. Chil. Hist. Nat., 13: 245-249.         [ Links ]

Rathbun, M.J. 1904. Descriptions of three new species of American crabs. Proc. Biol. Soc. Wash., 17: 161-162.         [ Links ]

Rathbun, M.J. 1907. South American Crustacea. Rev. Chil. Hist. Nat., 11: 45-50.         [ Links ]

Rathbun, M.J. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast. Proc. U.S. Nat. Mus., 38: 531-620.         [ Links ]

Rathbun, M.J. 1918. The grapsoid crabs of America. Bull. U.S. Nat. Mus., 97: 1-461.         [ Links ]

Rathbun, M.J. 1935. Preliminary descriptions of seven new species of oxystomatous and allied crabs. Proc. Biol. Soc. Wash., 48: 1-4.         [ Links ]

Rathke, H., 1837. Zur Fauna der Krym. Mem. Acad. Imp. Sci. St. Petersb., 3: 291-454, Plates 1-10.         [ Links ]

Renier S.A. 1804. Prospetto della classe dei vermi, nominati e ordinati secondo il sistema di bosc. Padua, 38: 15-27.         [ Links ]

Retamal, R.M. & L.A.Yáñez-Arancibia. 1972. Nuevo registro de Pinnixa bahamondei Garth, 1957 en Chile (Crustacea, Decapoda, Pinnotheridae). Bol. Soc. Biol. Concepción, 44: 103-107.         [ Links ]

Retamal, M.A. & H.I. Moyano. 2010. Zoogeography of Chilean marine and freshwater decapod crustaceans. Lat. Am. J. Aquat. Res., 38: 302-328.         [ Links ]

Retamal, M.A. & R. Trucco. 1973. Pinnixa valdiviensis Rathbun (Decapoda, Pinnotheridae) commensal de Urechis chilensis (Müller) (Echiuroida) - Registro de un nuevo hospedador. Bol. Soc. Biol. Concepción, 46: 75-79.         [ Links ]

Rioja, E. 1944. Estudios carcinológicos XVI. Observaciones sobre algunas especies de cangrejos del género Dissodactylus Smith (Braquiros, Pinnoteridos) de las costas mexicanas del Pacifico. An. Inst. Biol. México, 15: 147-160.         [ Links ]

Rodríguez, G. 1993. From Oviedo to Rathbun: the development of brachyuran crab taxonomy in the Neotropics (1535-1937). In: F. Truesdale (ed.). History of carcinology. Crustacean Issues 8. Rotterdam, Balkema, pp. 41-73.         [ Links ]

Runil-Ojeda, F. 2014. Caracterización genético-poblacional y variabilidad morfológica de Pinnaxodes chilensis (Decapoda: Pinnotheridae) en la costa oriental del Pacífico sur. Tesis de Magister, Universidad Austral de Chile, Valdivia, 114 pp.         [ Links ]

Sakai, T. 1939. Studies on the Crabs of Japan IV. Brachygnatha, Brachyrhyncha, 3. Yokendo Co., Tokyo, pp. 365-741, pls. 42-111.         [ Links ]

Sakai, T. 1969. Two new genera and twenty-two new species of crabs from Japan. Proc. Biol. Soc. Wash., 82: 243-280.         [ Links ]

Schmalfuss, H. 2003. World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beitr. Naturkd., Ser. A, 654: 1-341.         [ Links ]

Schmitt, W.L., J.C. McCain & E.S. Davidson. 1973. Fam. Pinnotheridae, Brachyura I: Decapoda I. In: H.-E. Gruner & L.B. Holthuis (eds.). Crustaceorum catalogus. W. Junk, Den Haag, 3, 160 pp.         [ Links ]

Semper, C. 1868. Reisen im Archipel der Philippinen. Zweiter Theil. Wissenschaftliche Resultate. Erster Band. Holothurien. Leipzig, Verlag von Wilhelm Engelmann, 177 pp.         [ Links ]

Silas, E.G. & K. Alagarswami. 1967. On an instance of parasitisation by the pea-crab (Pinnotheres sp.) on the backwater clam [Meretrix casta (Chemnitz)] from India, with a review of the work on the systematics, ecology, biology and ethology of pea crabs of the genus Pinnotheres Latreille. Symp. Ser. 2, Mar. Biol. Assoc. India, Part 3: 1161-1227.         [ Links ]

Smith, S.I. 1870. Notes on American Crustacea. Number I. Ocypodoidea. Trans. Conn. Acad. Arts Sci., 2: 113-176.         [ Links ]

Takeda, M. & P. Masahito. 2000. Systematic notes on the pinnotherid crabs of the genus Pinnaxodes (Crustacea: Decapoda: Brachyura). Bull. Nat. Sci. Mus., Tokyo, Ser. A (Zool.), 20: 99-112.         [ Links ]

Tesch, J.J. 1918. Goneplacidae and Pinnotheridae. The Decapoda Brachyura of the Siboga Expedition, 2. Siboga-Expeditie 39 (c1), livr. 84: 149-295, pls. 7-18.         [ Links ]

Torres, E.R. 2006. Primer registro para Argentina de Pinnixa valdiviensis Rathbun, 1907 (Decapoda, Pinnotheridae). Invest. Mar. Valparaíso, 34: 175-179.         [ Links ]

Walter, T.C. & G. Boxshall. 2015. World of Copepods database. []. Reviewed: 9 January 2017.         [ Links ]

Wells, G.P. 1963. The lugworms of the southern cold temperate zone (Arenicolidae, Polychaeta). Proc. Zool. Soc. London, 140(1): 121-159.         [ Links ]

Wells, H.W. & M.J. Wells. 1961. Observations on Pinnaxodes floridensis, a new species of pinnotherid crustacean commensal in holothurians. Bull. Mar. Sci. Gulf Caribb., 11: 267-279.         [ Links ]

White, A. 1843. Descriptions of apparently new species and varieties of insects and other Annulosa, principally from the collection in the British Museum. Ann. Mag. Nat. Hist., (12)1: 342-346.         [ Links ]

White, A. 1846. Notes on four new genera of Crustacea. Ann. Mag. Nat. Hist., [series 1], 18: 176-178.         [ Links ]

White, A. 1847. List of the specimens of Crustacea in the collection of the British Museum. British Museum, London, 143 pp.         [ Links ]

Williams, A.B. 1984. Shrimps, lobsters, and crabs of the Atlantic Coast of the Eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, DC., 550 pp.         [ Links ]


Received: 20 April 2016;
Accepted: 13 January 2017


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