Annotated type-catalogue of Brachyura (Crustacea, Decapoda) of the Muséum national d'Histoire naturelle, Paris. Part II. Gecarcinidae and Grapsidae (Thoracotremata, Grapsoidea), with an Appendix of pre-1900 collectors

ABSTRACT An annotated catalogue of the type specimens of two thoracotreme crab families, Gecarcinidae H. Milne Edwards, 1837 and Grapsidae MacLeay, 1838 (Crustacea, Decapoda, Brachyura, Thoracotremata, Grapsoidea) deposited in the collection of the Muséum national d'Histoire naturelle (MNHN), Paris, is provided. The Appendix lists the pre-1900 collectors, naturalists and donators mentioned in the text with the dates of expeditions or collecting trips and main countries visited. This provides valuable information on the sources of crustacean material in the MNHN historical collection. Neotypes are designated for the taxa: Gecarcinus lateralis Fréminville in Guérin-Méneville,1832; Grapsus cruentatus Latreille, 1803; Grapsus pictus Latreille, 1803; Grapsus varius Latreille, 1803. Lectotypes are designated for the taxa: Cardisoma urvillei H. Milne Edwards, 1853; Discoplax longipes A. Milne-Edwards, 1867; Gecarcinus lagostoma H. Milne Edwards, 1837; Pelocarcinus marchei A. Milne-Edwards, 1890; Thelphusa rotunda Quoy & Gaimard, 1824; Grapsus brevipes H. Milne Edwards, 1853; Grapsus fourmanoiri Crosnier, 1965; Grapsus gracilipes H. Milne Edwards, 1853; Grapsus granulosus H. Milne Edwards, 1853; Grapsus grayi H. Milne Edwards, 1853; Grapsus lividus H. Milne Edwards, 1837; Grapsus maculatus H. Milne Edwards, 1853; Grapsus maurus Lucas, 1846; Grapsus oceanicus H. Jacquinot in Hombron & H. Jacquinot, 1846; Grapsus ornatus H. Milne Edwards, 1853; Grapsus peroni H. Milne Edwards, 1853; Grapsus pharaonis H. Milne Edwards, 1853; Grapsus plicatus H. Milne Edwards, 1837; Grapsus webbi H. Milne Edwards, 1853; Leptograpsus ansoni H. Milne Edwards, 1853; Leptograpsus bertheloti H. Milne Edwards, 1853; Leptograpsus gayi H. Milne Edwards, 1853; Leptograpsus rugulosus H. Milne Edwards, 1853; Leptograpsus verreauxi H. Milne Edwards, 1853; Metopograpsus eydouxi H. Milne Edwards, 1853; Metopograpsus gracilis Saussure, 1858; Metopograpsus intermedius H. Milne Edwards, 1853; Metopograpsus maculatus H. Milne Edwards, 1853; Metopograpsus miniatus Saussure, 1858; Metopograpsus pictus A. Milne-Edwards, 1867; Pachygrapsus minutus A. Milne-Edwards, 1873; Pachygrapsus striatus A. Milne-Edwards, 1873.


INTRODUCTION
In response to the urgency to inventory the world's fauna and provide solid, reliable support for taxonomic research and for the knowledge of the biodiversity, an essential task is to accurately identify the specimens upon which taxa have been established, i.e., the types that are name-bearing or onomatophores (Simpson 1940;Dubois 2000Dubois , 2005Dubois & Nemésio 2007;Dubois et al. 2014). The purpose of this work is to clarify the taxonomic status of the nominal taxa, the nomen of a taxon being attached to a type specimen and type locality as stated by the International Code of Zoological Nomenclature (ICZN 1999, Chapter 16), referred to as the ''Code'' throughout this text. The Code (Recommendation 72F.4) recommends that every institution publishes "lists of name-bearing types in its possession or custody", following strict rules of typification. This requires that experts explore on which original specimens the binominal scientific names were based since the Systema Naturae of Linnaeus (1758).
The practice of clearly highlighting the type material was not followed until relatively recent times when implementation and special attention were given to particular individuals or samples that were supposed or suspected to represent type material. This applies mainly to the dry and historical collection of Crustaceans and to early papers at a period where the designation of types was not in use. In these cases, identification of the type specimens is particularly difficult, which may lead to incorrect designation and misapplication of the Code.
There may be several major problems with historical collections preserved in dry condition. One significant obstacle to unequivocal interpretation is the fact that the original collectors and authors did not label the specimens included (i.e., original, handwritten labels or inscriptions have not survived, apart from exceptional cases). This contributes to uncertainty regarding typification issues (see Material and Methods).
The Code (Art. 72.4.1) states that "The type series of a nominal species-group taxon consists of all the specimens included by the author in the new nominal taxon", thus our first step was to identify this material. The most crucial task was to decide which sample, whether labelled as "type", "cotype" or "syntype", or not labelled as such, actually represents the material upon which the original author based the description of a new nominal species and thus representing the type or belonging to the type series. The decision that a particular specimen is actually the original one used in the description of the species must be made after detailed historical and scientific investigations of the material in question. This critical examination is finally based on the labels of subsequent workers (researchers and curators). In the absence of any explicit designation, the selection of the type specimen is based on a presumption ("presumed" type). Although impossible to state, evidence may be obtained by carefully examining the labels, specimen presentation (glue, wire-or wood-stick remnants), or using reliable sources, i.e., the locality, name of the collector, date of the supposed collection, and the original publication in which measurements and figures were possibly provided, and additionally a review of the relevant literature.
According to the Code (Art. 72.4.1.1; 73.1.1; 74.6), for a nominal species or subspecies established before 2000, any evidence, published or unpublished, may be taken into account to determine which specimens constitute the type series. If the nominal species-group taxon was based on a single specimen, either so stated or implied in the original publication, that specimen is the holotype fixed by monotypy. If the number of specimens was not clearly specified in the original publication, with the description perhaps based on several specimens, a lectotype is selected as unique bearer of the name of the nominal species-group taxon, whether only one specimen or more than one has been found in the collection; the remaining syntype(s) become(s) paralectopype(s).
Fixation of lectotype by inference of "holotype" or "type" is a particular case where the first author to have published (before 2000) the assumption that the species was based upon a single type specimen (holotype by monotypy), instead of being based on more than one specimen or without mention of the number of examined specimens, is deemed to have designated that specimen as the lectotype (Code, Art. 74.6). In cases where it was impossible to determine with certainty whether any of the extant specimens from the type locality in the collection corresponds to the type material, the holotype is regarded as lost, and a neotype must be designated. Conversely, in cases where specimens indicated as types have been prove to be incorrectly labelled or published as types, a well-argued correction or reappraisal is necessary.
Further investigations sometimes show that specimen(s) presumed to be the single known presumed holotype or syntypes was (were) not the only one(s) belonging to the type series. It may be found that one or more samples existed, being deposited in another institution, a current data not yet recorded. Some subsequent type designations, largely overlooked, have thus been already made through gifts or exchanges of original material, of syntypes. For example, some syntypes donated to the RMNH or exchanged through H. Milne Edwards and V. Audouin led to the publication of new names by W. de Haan in the Fauna Japonica (see Fransen et al. 1997). Duplicates were also later exchanged by A. Milne-Edwards with the RMNH. See other examples in Cleva et al. (2007: 6).
An important source for the identification of the type series material is the name of the collector that sometimes appears on the "original label" (see Material and Methods) and may give a valuable indication of the period when the specimen(s) was(were) collected. The Appendix included herein provides information on the voyager-naturalists, collectors and donators mentioned in the text, the dates of their voyages or collecting trips and the main countries visited. This provides useful information on sources of crustacean material in the Muséum national d'Histoire naturelle (here referred to as "the Muséum", or MNHN) historical collection. This list is restricted to those persons that contributed specimens in the two families studied here, but will be useful to all taxonomists. The same collectors supplied material not only in all other brachyuran families, but also of all kinds of animals and plants collected during their voyages.
Another precious source of information is the "Catalogue of Articulated animals: Crustaceans, Arachnids, Insects, received, donated, exchanged or bought" ("Catalogue des Ani maux articulés : Crustacés, Arachnides, Insectes, reçus, donnés, échangés ou achetés") (called CAA). It consists of a series of handwritten hard-bound books dating from 1826. Deposited in the Entomology Collection, it records the entry date of the arthropods in the MNHN and includes various data, such as date and number of the entry, geographical origin, collector, donator, exchanges, identification, and sometimes number of specimens and other details (Fig. 1A). It was developed at a time when crustaceans and insects (and also worms) belonged to the same chair. It represents one of the oldest catalogues dealing with the Crustacea housed in the MNHN. The two CAA numbers for each sample correspond to the entry number and year of entry into the MNHN, respectively: e.g. for 225-63 the second number designates 1863.
Two other historical catalogues are those handwritten by P. A. Latreille himself, with bookbindings dated 1807 and 1814. They are deposited in the Entomology Library and herein designated as "LC1807" and "LC1814" (Cleva et al. 2007: figs 3, 4). The parts concerning the Crustacea collection list the content of the various "cadres" that referred to the showcases of the Muséum galleries at that time.
Furthermore, hard-copy registers listing the entirety of the crustacean material are deposited in the Crustacea section of the "Direction générale déléguée aux Collections" [ The MNHN has started an inventory of all its contents, the collation of all data and the registration of the entire collection, with a new numbering system (see Material and methods). One of the ultimate purposes is the typification of all the species.
The first annotated catalogue of brachyuran type specimens preserved in the MNHN collection was carried out on the section Podotremata Guinot, 1977 (crabs with male and female coxal gonopores, and spermathecae) and published by Cleva et al. (2007). Therein, the types of 104 nominal species are listed and photographed. Experience has shown that an important part of the surviving brachyuran material collected during the numerous voyages and renowned expeditions of the French naturalists and travellers ("voyagers-naturalists", see Laissus 1981;Bauchot et al. 1990Bauchot et al. , 1997 from the mid-18 th and 19 th centuries has been preserved until today. The present catalogue focuses on the brachyuran namebearing type specimens of the section Thoracotremata Guinot, 1977 (crabs with male and female sternal gonopores). In an initial phase, the types of nominal species of two families of Grapsoidea MacLeay, 1838, the Gecarcinidae H. Milne-Edwards, 1837 andGrapsidae MacLeay, 1838, are listed. Identification and allocation of the type material are fundamental today for the objective identification of names. In order to preserve stability of nomenclature there is a need to fix the status of species for comprehensive taxonomic revision and phylogenetic analyses of family-group taxa. Since the development of molecular techniques has shown the exis tence of cryptic species that far exceed earlier expectations using traditional approaches, the selection of a neotype, even for presently non-valid species, is required to stabilize the use of the names.

MATERIAL AND METHODS
The present paper follows Cleva et al. (2007), with only minor modifications. The names of the taxa are presented here in their original combination; genera and species are listed in alphabetical order for each family. Type specimens of currently invalid species (as junior synonyms, junior homonyms, unjustified emendations, unnecessary substitute names, or suppressed names) are also listed. The current status is specified, as far as known. This annotated catalogue has benefited from the contribution of three scientists who were able to verify the identifications, the conformity between the labels and crabs, and the current status of species. The authors are well versed to check and discuss nomenclatural problems.
The original published description of each species is cited. The publication in which holotype, lectotype or neotype have been selected is likewise provided. No synonyms are provided. Other information is placed in the remarks, based on official published sources, often with the citation of the most recent reference providing a complete description of the type(s).
The occurrence of multiple labels for a single sample is very frequent, most labels not being signed. They are all cited in our text, as "Original label" and "Additional label(s)". In the case of the historical dry collection, there is practically never an available label handwritten by the author; thus, what we refer to as the "original label" is what we suppose to be the first label, with all the data transcribed verbatim in the original script (generally in French), and in writing in italics all the taxa names. The historical boxes had either glass sides supported by a metallic frame or were constructed of orange cardboard sides fixed to a glass top, with the crabs glued on to a white cardboard by their ventral surface, often with a piece of elder (Cleva et al. 2007: fig. 1A, B). They contained the "original data" meticulously calligraphed in black ink italics ("fixed labels"), probably produced during an extensive curatorial reorganisation and reconditioning of the whole collection, the date of which is as yet unknown. The geographical name that is written on the label is cited as in use when the specimen was collected. In the past, the provenance was frequently a country rather than a precise locality, often without indication of the collector and without any date. The mention "Coll. A. Milne Edw. 1903" on the labels of some historical dry or in alcohol specimens does not mean that the material was collected by A. Milne-Edwards in 1903 but must be interpreted as "Collection of A. Milne-Edwards, 1903", indicating, to the best of our knowledge, that the material was included in the MNHN collection at this date, perhaps during its reorganisation. These old boxes have been progressively replaced by easier-to-open, all-plastic boxes; the specimens were separated from the old glue, most being now fixed onto a corkboard by pins (Cleva et al. 2007: fig. 1C, D). The old cardboard was either kept to ensure safeguarding of the information or, when impossible, the data were carefully rewritten, becoming the "original label". After the collection was partly flooded in 1991, a long process of restoration of damaged specimens began in 1993, and a part of the dry historical collection of brachyurans has been treated with acetone and rhodopas. After almost two centuries (and surviving world wars and natural disasters), the dry material of the MNHN historical Crustaceans collection can be considered well preserved, with specimens being generally complete and in good condition, together with their corresponding labels. Special care is taken to maintain it in good condition for other several hundred years.
A new registration method has been implemented in the early 2000s for all MNHN collections in order to build a collective database. A unique alphanumeric registration number is attributed to each sample; for the Crustacean collection, this new registration number consists of MNHN follow by IU (for Invertebrates, Crustacea), a year and an incrementing number, i.e., "MNHN-IU-20XX-XXXX. The old registration number for brachyuran crabs consisted of MNHN followed by B (for Brachyura) and an incremented number, i.e., "MNHN-BXXXX". Presently, both the new and the old registration numbers are strictly connected and both are valid, e.g. MNHN-IU-2014-11214 (= MNHN-B26950). All the registered material is available online (https://science.mnhn. fr). Internet access to this material will further increase the value of the collection and benefit researchers and naturalists worldwide.
When a lectotype is designated from the syntype series in one sample, it is separated with the original registration number, and a new registration number is assigned to the remaining individuals of the sample.
The condition of samples is noted as "Preservation": dry or in alcohol (= ethanol 75%), completely or variably damaged; the absence of information means that the specimen(s) is (are) in a good state of conservation and whole. A few dry specimens have been rehydrated, sometimes only partly (gonopods), for research purposes.
When the type material is not held by the MNHN or if a part is known to be deposited elsewhere, the institution where it is preserved is indicated. Information on type data and depository institutions have been carefully provided by Davie (2002: 11), an asterisk (*) meaning this has not been verified by him, and the statement 'probable depository institution' meaning uncertainty: some data proved to be accurate and have only been supplemented here; others required correction.
A number of original specimens supposedly preserved in the MNHN collection, i.e., those that are recorded in the original publication or subsequent literature as being described by French scientists such as Lamarck, Latreille, Guérin, H. Milne Edwards and A. Milne-Edwards, could not been traced at the present time. They are not included in this catalogue. A listing of all the species that might be expected to be represented in MNHN collections could lead to hazardous or erroneous results, the discovery of forgotten specimens always being possible. One such case is the type material of two Gecarcinidae (Cardisoma hirtipes Dana, 1851 and Cardisoma obesum Dana, 1851) which were recently found in old boxes in the dried collection of the U.S. National Museum of Natural History (USNM), Smithsonian Institution (Ng 2017).
The types of 42 species cited herein are deposited at the MNHN, with one exception: Grapsus albolineatus Lamarck, 1818. The type material of this species is no longer extant in our Collection but, since the specific name raises a nomenclatural problem, a discussion about this controversial topic is included.
Most type specimens are illustrated (Figs 2-7, photographs by N. Mollaret). Measurements are reported in millimetres (mm) as carapace length (cl) × width (cw), taken at its maximum (including teeth or spines if present).    Alphonse Milne-Edwards (1873a: 294) regarded Disco plax longipes as "very rare" in New Caledonia. The specimen was simply labelled as from "New Caledonia", without any other indication as is often the case in old descriptions and records, but the presence of the species on the main island of New Caledonia is doubtful. It seems that, since A. Milne-Edwards (1867), no specimens of D. longipes have been collected from the main Island where the species is not known locally (Ng & Guinot 2001 Rathbun 1918: 361, under Gecarcinus lagostoma;1918: 365, under ?Gecarcoidea lalandii;Türkay 1973: 96). Thus, the locality "Australasia" of the lectotype and paralectotypes must be recognised as inaccurate, and Brazil must be considered the type locality of Gecarcinus lagostoma, whereas "Australasia" is the original type locality of Gecarcoidea lalandii. Lai et al. (2017: 410, 416 Hobbs 1969;Türkay 1970Türkay , 1974aBliss et al. 1978;Hartnoll 1988;Ng & Guinot 2001;Ng et al. 2008;Perger & Wall 2014). Low et al. (2013) stated that Guérin (1832 in Guérin 1829-1837) must take precedence. Fréminville (1835) described Ocypoda lateralis from La Désirade, Guadeloupe, Marie-Galante, Martinique, Les Saintes in the West Indies, based on specimens that he has probably himself collected (see Appendix). In the introduction of this paper (1835: 213, footnote), the editor of the volume, actually H. Milne Edwards, indicates about Fréminville's "tourlourou": "son Ocypoda lateralis est une espèce de Gécarcin bien distincte et dont j'ai donné une figure dans mon Histoire naturelle des Crustacés. M. Guérin l'a également figuré dans son Icono graphie du règne animal" ["his Ocypoda lateralis is a clearly distinct species of 'Gécarcin' and to which I have given a figure in my Histoire naturelle des Crustacés. M. Guérin has also figured it in his Iconographie du règne animal"]. Guérin (1832, in Guérin 1829-1837: 7) had previously used the name "Gecarcinus lateralis Fréminville" and illustrated what is clearly the same species. It is very likely that Guérin used the same material as Fréminville and that he was aware of the future species by Fréminville. According to the official dates, Guérin's plate has been published before the description by Fréminville in 1835. According to Low et al. (2013) the authorship of G. lateralis must be credited to Guérin (1832). But, as Fréminville is solely responsible for the species name in a way satisfying the criteria of availability (Code, Art. 50.1.1), the authorship of G. lateralis should be credited to Fréminville in Guérin (1832).
Türkay (1970), based on erroneous indications given by J. Forest, the curator of Crustacea Collection in the MNHN at the time, regarded as the type series of G. lateralis material collected by two French naturalists, Beaupertuis and Bélanger, and deposited in this institution. Türkay (1970: 337) selected as lectotype of Gecarcinus (Gecarcinus) lateralis a specimen from Guadeloupe collected by Beaupertuis. Türkay Bélanger's specimens are likely to not have been collected during this period (i.e., before 1832), these two naturalists having traveled in the Antilles later on as shown by the entry dates of their material in the CAA (Beaupertuis: 1839 from Guadeloupe; Bélanger: 1859, 1860, 1864 from Martinique) (see Appendix), and cannot constitute the original material seen by Guérin (1832).
According to the Code (Art. 74.2), "the name-bearing type of any nominal taxon, once fixed in conformity with the provisions of the Code, is not subject to change" except if the lectotype is "found not to have been a syntype". Thus, "if it is demonstrated that a specimen designated as a lectotype was not a syntype, it loses its status of lectotype". A neotype of G. lateralis must be fixed. Having been previously referred to, although unsupported, as the lectotype for the species by Türkay (1970), then by Perger & Wall (2014), and considering its locality, the specimen from Guadeloupe registered MNHN-IU-2000-3758 (= MNHN-B3758) is eligible for neotype designation: it is hereby selected as the neotype of Gecarcinus lateralis Guérin, 1832. . Quoy & Gaimard (1824: 527) did not explicitly mention if the taxon was based on a single individual, but their description provides information on the behaviour of several crabs: "ces animaux extrêmement défiants" ["these extremely challenging animals "], an indication that numerous specimens were probably collected in Guam. "Océanie" is a large area that can include Guam, thus the MNHN specimen such labelled may constitute a part of the type material. Türkay (1974a: 971, 972; pers comm., Jan. 2013) did not considered any specimen as type of Thelphusa rotunda, assuming the species was a synonym of Cardisoma frontalis H. Milne Edwards, 1853, and regarded the specimen of Quoy & Gaimard (1824)  remArks The binomen "Grapse albolineatus" ("grapse rayé de blanc" or white-striped grapse) has been introduced by Latreille, in Milbert (1812: 275) but Grapsus albolineatus is usually credited to Lamarck (1818: 249, as Grapsus albolineatus, "grapse raies-blanches"). As is often the case, Latreille in Milbert (1812) and Lamarck (1818) were based on the same material, namely a crab collected by Mathieu from Mauritius (= Île de France; see also Latreille 1825: 148), and referred to the same species. Latreille in Milbert (1812: 275) wrote "grapse" ("Ce sont les grapses de M. Lamarck" [These are the grapses of M. Lamarck]). Although "G. albolineatus" is a binominal combination with latinised species names, the absence of any diagnosis makes it not available (see the discussion below). The genus name Grapsus belongs to Lamarck (1801: 150), with Cancer grapsus Linnaeus, 1758 as type species by tautonomy (see also designation by Latreille 1810: 422).
In most French and German scientific works at this time, the Latin name was accompanied by its translation in French (e.g. Latreille 1806; Lamarck 1818; H. Milne Edwards 1834, 1837), or in German, (e.g. Herbst 1782-1804). These specific spellings are only the latinisation of the names depicting morphological (shape, colour, ornamentation), behavioural, geographical and other features, by simple juxtaposition of both names. This simple translation of the names is not enough to constitute a diagnosis and to make the taxa available, and does not correspond to a brief description. These names cannot satisfy the requirement of Article 12.1 of the Code stipulating that (for the names published before 1931) to be available a taxon must be accompanied by a description (even if only the colour) or a definition of the taxon that it denotes, or by an indication. According to Ng et al. (2008: 22) these three species were "defined by their colour or shape", but this is not the case and these names are not available. Moreover, Ng et al. (2008: 22) have treated the three above-mentioned grapses in the same manner as the case of Grapsus tessellatus, and credited Grapsus albolineatus, Grapsus erytrhocheles and Grapsus tuberculatus, like G. tessellatus, to Latreille in Milbert (1812). It is an erroneous generalisation: G. tessellatus is an available name with the authorship of Latreille in Milbert (1812), whereas the other three names are clearly nomina nuda. Grapsus albolineatus belongs to Lamarck, 1818, G. erytrhocheles is not available (or at least corresponds to a species inquirenda); G. tuberculatus must be credited to Lamarck (1818: 247, as Plagusia tuberculata) (Schubart & Ng 2000: 327, 334, fig. 3A).
To avoid ambiguity, a point that requires discussion is the meaning of "vernacular". According to the English glossary of the Code (ICZN 1999: 110) a vernacular name is "A name of an animal or animals in a language used for general purposes as opposed to a name proposed only for zoological nomenclature", whereas a zoological name is "The scientific name of an animal taxon in binominal nomenclature". Thus in French scientific works at the period of Henri Milne Edwards (1834, 1837), the new taxa are proposed in a scientific context and are not vernacular names, conversely to the views of most carcinologists (including Ng et al. 2008: 20).
The type material from Mauritius collected by Mathieu was not found in the MNHN collection. A neotype should be fixed for Grapsus albolineatus, preferably from the type locality, Mauritius or its proximity (Code, Art. 75.3.6), as it is considered the senior synonym of numerous species (see Banerjee 1960: 147, 154;Holthuis 1977: 145, 147), such as G. strigosus (Herbst, 1799), from the Indian Ocean; G. (Go niopsis) flavipes MacLeay, 1838, from Cape of Good Hope; G. peroni H. Milne Edwards, 1853, from Australia (see below); and G. longipes Stimpson, 1858, from Kikaishima, southern Japan, and Hong Kong (Davie 2002). The type locality " les mers de l'Ile-de-France" and the occurrence of this species on the east coast of Africa was questioned by Crosnier (1965: 17) but according to Holthuis (1977: 147) it is one on the most common and conspicuous crabs of the Red Sea.
A neotype designation is unfortunately not possible at this time because the MNHN collection does not possess any material from Mauritius or a nearby locality (Code, Art. 75.3.6). Grapsus

remArks
The description did not mention how many specimens were examined to diagnose the species. Only one specimen was found in the MNHN collection but, as there is no assumption it is a holotype (i.e., fixed by monotypy), it is designated here as lectotype (Code, Recommendation 73F). remArks Latreille (1803: 70) described this species from the islands of South America ("îles de l'Amérique méridionale"), imprecise information (as it was common in the past) indicating the geographical origin. Henri Milne Edwards (1837: 85) has added two localities, Brazil and Antilles, based on specimens present at that time in the collection of the MNHN (as "C.M."). Today, the MNHN collection contains several samples of dry specimens labelled Grapsus or Goniopsis cruentatus, from Brazil, Mexico, French Guiana and Martinique. They were all collected too late to belong to the type series; moreover, some of the localities cannot be considered "islands of South America". For example:

Grapsus cruentatus
(1) the sample with one specimen MNHN-IU-2000-3406 (= MNHN-B3406) collected from Brazil by Delalande in 1816 (see Appendix), a date that is too late to have enabled Latreille (1803)  Concerning the sample from Martinique in the MNHN collection, which could correspond to the type locality, consisting of two males collected by M. Plée, it cannot belong to the type series: the material of Plée was collected after 1820 (Bauchot et al. 1990: 117;see Appendix), and this is confirmed by the entry date "1826" of Plée's material in the CAA. Since no MNHN material conforms to a presumed syntype and no name-bearing type is believed to be extant, a neotype should be designated. The more complete male of the MNHN-B3406 sample has been selected as neotype, MNHN-IU-2000-3407 (= MNHN-B3407). The slightly smaller male specimen has been separated and is registered as MNHN-IU-2000-1109 (= MNHN-B3407).    fig. 1) and Holthuis (1977: 147, 148) commented that it is a distinct species (see also Vannini & Valmori 1981;Zaouali et al. 2007        In the handwritten Catalogues of Crustacea by Latreille (LC1807, LC1814), two specimens, a male and a female, are cited without locality, date or collector: there is no mention of bopyrids (Fig. 1B). The figure of Grapsus pictus in Latreille (1803: pl. 47, fig. 2) does not show any deformation of the carapace by a parasite but such an unnatural shape was perhaps not taken into account by the drawer. Even if Latreille (1803) could have confused the preadult female with a male, we do not have enough objective proof to consider that these specimens belong to the type series of Grapsus pictus. Consequently, for clarification purposes the designation of a neotype is justified. The MNHN collection contains one specimen, a male labelled "Grapsus rudis, Sandwich Islands, Quoy & Gaimard" with the mention "Type" on the back of the cardboard box (MNHN-B10875). Castro (2011: 114) in his catalogue of Brachyura from the Hawaiian Islands regarded it as the holotype of G. rudis, actually a holotype by monotypy.
Our examination of the specimen MNHN-IU-2000-10944 (= MNHN-B10944) has confirmed that Leptograpsus berth eloti is indeed Pachygrapsus marmoratus, and not Leptograpsus variegatus (Fabricius, 1793) as stated by Kingsley (1980). Recent publication on the phylogeography of Pachygrapsus marmoratus from the African Mediterranean coast (Deli et al. 2016) has revealed a genetic homogeneity across the Siculo-Tunisian Strait.  remArks Rathbun (1918: 245) mentioned that the type was in "Paris Museum" but without listing any material. Although Lep tograpsus rugulosus was recognised by Kingsley (1880), it is the junior synonym of Pachygrapsus transversus (Gibbes, 1850) according to Poupin et al. (2005: 44). The MNHN collection contains two samples that could have been seen by H. Milne Edwards in 1853: a complete and dry female specimen MNHN-IU-2000-3535 (= MNHN-B3535), collected from Rio de Janeiro by Peter Claussen (see Appendix), and a specimen MNHN-IU-2000-3534 (= MNHN-B3534) from Brasil collected by Charles Gaudichaud-Beaupré, who was a well-known, extremely active collector at that time in the MNHN (see Appendix). The first entry in the MNHN of South American material collected by Gaudichaud is from 1832, as indicated in the CAA. Henri Milne Edwards was used to mention the localities in accordance with the information provided by the collector, thus the type locality is "Brésil" and not "Rio de Janeiro". The specimen MNHN-IU-2000-3534 (= MNHN-B3534) of Gaudichaud is selected as lectotype. preservAtion. -Carapace cracked on right side at cardiac region, RP4 missing.
In assuming a holotype rather than designating a lectotype, Türkay (1974b: 145) proceeded as though syntypes did not exist. But there is a specimen, a dry female, collected by Saussure from the same locality, St Thomas, deposited in the MNHN collection, MNHN-IU-2000-10991 (= MNHN-B10991), as confirmed by the mention in the CAA of an exchange between Saussure and the MNHN in 1858, entry 270, corresponding to "270.58" on the original label. We may consider that this specimen has been collected at the same time as the holotype and could belong to the syntype series. Therefore, Türkay (1974c), who has published the assumption that the species-group taxon was based upon a single type specimen, is deemed to have designated that specimen as the lectotype (Code, Art. 74.6: Fixation of lectotype by inference of "holotype"). The female MNHN-IU-2000-10991 (= MNHN-B10991) may be considered paralectotype.  (1949: 470), Gordon found two specimens in the MNHN labelled as "cotypes", patria ignota, but her sketches of the G1 of each of these two "cotypes" revealed that they belonged to two different species. Tweedie (1954: 127, 128, under M. gracilipes fig. 17) indicates the presence of a parasite deforming the carapace. In assuming a holotype rather than selecting a lectotype, Türkay (1974c: 142, 143) proceeded as though other syntypes did not existed. But we have found supplementary material, a male MNHN-IU-2000-10994 (= MNHN-B10994), collected by Saussure from St Thomas, presumably at the same time, as confirmed by the mention in the CAA of an exchange between Saussure and the MNHN in 1858, corresponding to the entry "262.58". This specimen likewise shows a deformation of the carapace. The fact that both specimens are similarly parasitised may be an indication that they should be from the same lot and, presumably, they belong to the syntype series, despite there being no indication of swelling on the carapace in Saussure's paper. It is reasonable to expect that the original description was based on more than one specimen. Therefore, Türkay (1974c), who published the assumption that the species-group taxon was based upon a single type specimen in recognising a holotype, is deemed to have designated that specimen as the lectotype (Code, Art 74.6). The male MNHN-IU-2000-10994 (= MNHN-B10994) may be considered a paralectotype. It is also the case with Metopograpsus gracilis Saussure, 1858, also from Saint Thomas (see above).      MNHN) for access to the collection. We are very grateful to the referees for their careful reading and constructive comments on the manuscript. We deeply thank Annemarie Ohler for providing her experienced advice on nomenclatural matters. Many thanks are due to Claudia Ratti for her English text revision.

Metopograpsus pictus
The project started with the previous curator, Regis Cleva, after the publication of Part I of the Catalogue of brachyuran types deposited in the Muséum national d'Histoire naturelle, dedicated to Podotremata (Cleva et al. 2007). His invaluable help for the compilation of the type material as well as his support to the first author at the beginning of the project are acknowledged.
The authors would like to thank e-ReColNat (ANR-11-IN-BS-0004) for the use of the numerous photos of the type materials deposited in the MNHN, the team of the MNHN Entomology Collection (L. Albenga and A. Mantilleri) for access to the CAA, and the MNHN Libraries, especially J. Guglielmi, librarian in the MNHN Entomology section for access and photography ( Fig. 1) (2010). Another collector named Beaudoin, also known as a captain, deposited in the MNHN numerous crabs, often in dry condition, collected from the Red Sea and the Mediterranean. This is significant for knowledge of Egyptian carcinology: they can be also traced in the CAA, with 1866 as the entry date (Guinot & Cleva 2009: 28). (1807-1871). Also known as Luis Daniel Beauperthuy, often indicated as "Beaupertuis" on the labels, was born in Guadeloupe and died in Guiana. Since childhood, he was in contact with nature. In 1838, he was sent as a naturalist voyager by the Muséum d'histoire naturelle at Paris, to Venezuela where he collected a huge number of specimens (reptiles, fishes, molluscs, crustaceans, plants, etc.), now deposited at the MNHN. In 1853 he discovered that yellow fever is transmitted by mosquitoes and not through the air. (Bauchot et al. 1990(Bauchot et al. , 1997. (1805-1881). He was a French explorer and naturalist. In 1825, he was commissioned to create the botanical garden in Pondicherry, India, where he arrived in 1826 after a fourteen-month voyage! Along the way, he collected many documents and valuable ethnographic material. Bélanger made three more trips, especially one in Java, Indonesia. In 1850, he was appointed as the director of the botanical garden in Martinique wich rapidly turned into a scientifically important institute: the garden quickly became a valuable storehouse of many rare plants and the only one in the French colonies. During his stay in Martinique, Bélanger continued to enrich the collections of the MNHN (Broc 1992;Bauchot et al. 1990Bauchot et al. , 1997. In the CAA, entries of material collected by Bélanger as "botaniste du roi à Pondichéry" ["the king's botanist at Pondicherry"], are mentioned for 1828 without origin, then for 1859, 1860 and 1864 from Martinique.

Dumont d'Urville, Jules sébastien césar (1790-1842).
He was an explorer, cartographer, botanist, linguist and writer. In 1807, at the age of 17, he joined the navy. In 1820, he was sent on a voyage to the eastern Mediterranean Sea on board the Chevrette. In 1822, he set off on the first of his scientific discovery voyages with the Coquille, which sailed from Toulon, with Louis Isidore Duperrey as commanding officer and Dumont d'Urville as second in command. The voyage lasted 31 months. Dumont d'Urville visited New Zealand for the first time in 1824. In 1826, after being promoted to the rank of commanding officer, he sailed from Toulon on another voyage that took him to New Zealand. His ship, the Coquille, was rechristened as Astrolabe in honour of one of La Pérouse's ships. Many important botanical and entomological investigations were carried out during that voyage by Dumont d'Urville, Jean René Constant Quay, Joseph Paul Gaimard (see under these names) and Pierre Adolphe Lesson. The Astrolabe arrived back in Marseille in 1829 where the precious cargo of specimens was unloaded before the final docking in Toulon. Dumont d'Urville was commanded to publish an account of the Astrolabe's voyage, which comprises 12 volumes and five albums; it was completed in 1835. In 1837, Dumont d'Urville left Toulon for his third voyage to the southern oceans. He had two ships, the Astrolabe, which he commanded personally, and the Zélée, under the command of Captain Charles berthelot, sabin (1794-1880). A French naturalist and ethnologist, who resided in the Canary Islands for part of his life. After the Napoleonic Wars, he joined the merchant fleet, travelling between Marseille and the Antilles. He first visited the Canary Islands in 1820, which he considered a paradise for naturalists. He taught in Tenerife, managed the botanical gardens in Orotava and studied the natural history of the Canary Islands. He was joined by Philip Barker Webb (see under this name) in 1828, and by 1830 they had collected sufficient information to publish the first volume of L'Histoire Naturelle des Îles Canaries [The Natural History of the Canary Islands] in Geneva in 1835. Berthelot focussed on the ethnography, history and geography of the islands, while Webb completed the natural history sections (Mearns & Mearns 1988;Bauchot et al. 1990Bauchot et al. , 1997. (1866-1938). Initially a musician, he was a traveler and historian of Polynesia. After lessons in natural history by the Muséum in 1894, he was appointed to collect natural history material. He travelled extensively around the world, in Australia, New Zealand, South America and finally Polynesia to which he devoted the main part of his life, producing two volumes to its civilisation. In 1912-1914, he embarked on a trip to Oceania. He returned to Paris in July 1914, with many documents, and published several contributions dedicated to the history, traditions, religions and culture of Polynesians (Aimès 1952). (1804-c. 1855). He was a Danish natural history collector, born in Copenhagen, but obliged to leave his country, joining the Brazilian army. He bought a farm in Minas Gerais, where he established a veritable museum and maintained natural history collections. He collected numerous plants, fossils and mineral specimens and sold them to the British Museum and the MNHN (Bauchot et al. 1990, 1997.

claussen, Peter
Delalande, also known as lalande, Pierre antoine de (1787-1823). French naturalist and explorer. After studying painting, at the age of 13 he entered the Muséum d'histoire naturelle, where his father was a taxidermist. As an assistant naturalist to E. Geoffroy Saint-Hilaire, he accompanied Auguste de Saint-Hilaire (see under this name) on a trip to Brazil in 1816. Here they collected samples of many animal species in the region of Rio de Janeiro. In April 1816, Auguste de Saint-Hilaire and Pierre-Antoine de Lalande left France aboard the frigate L'Hermione towards Brazil but in December 1816, for health reasons, Delalande returned to France carrying the specimens. In 1818, he began an expedition to South Africa with his nephew Jules Verreaux who was around 12 years old. They travelled and collected natural history material in South Africa for three years. On their return in 1821, they brought an astounding number (131 405) of specimens, mostly plant material, but including 288 mammals, 2205 birds, 322 reptiles, 265 fish, 3875 shellfish, and various human skulls and skeletons. After the Restoration (1814-1830) he changed his name from Lalande to Delalande. His nephew (see under Hector Jacquinot (see under this name). He received a royal command from King Louis-Philippe to publish an account of the voyage. Many islands and places have been named in honour of his valuable chartings (Bauchot et al. 1990(Bauchot et al. , 1997Clark & Crosnier 2000;Dunmore 2007;Duyker 2014). See also under Hombron, Jacques Bernard. Fortuné Théodore (1802-1841). A French naturalist, Doctor of Medicine, he was surgeon and natura list on the corvette La Favorite that made a circumnavigation in 1830-1832 captained by Cyrille Pierre Théodore Laplace. In 1836-1837, with Louis François Auguste Souleyet (see under this name), he voyaged on La Bonite captained by Auguste Nicolas Vaillant. La Bonite sailed from Toulon and visited many places, including Rio de Janeiro, Valparaiso, Peru, the Sandwich Islands, the Philippines, Macao, Vietnam, Singapore, India, Réunion Island and the Cape. The collections brought back surpassed the hopes of the professors at the Muséum. Eydoux was nominated "correspondant du Muséum" in 1840, to which he donated his numerous collections. When he was just beginning to write the zoological part of this journey, Eydoux was sent to Martinique where he died of yellow fever. The results of the collected material were published with Louis Souleyet (see under this name) in the Voyage autour du monde […] sur la corvette La Bonite [Voyage around the world […] on the corvette La Bonite], and, after Eydoux's death, Souleyet continued its publication, with Eydoux as co-author. The publication comprises 18 volumes and three atlases, from 1840 to 1866(Bauchot et al. 1990, 1997.

Fontaine, P. a.
Little is known about him except that several natural history specimens brought to the Muséum by Alcide d'Orbigny from his voyage of exploration in South America (1826-1834), were collected by Fontaine in Chile and Peru and that he was of a great help for d'Orbigny. Many crab species are labelled with his name, either on its own or along with those of Gay, d'Orbigny, or Gaudichaud (Guinot & Cleva 2002). Several invertebrate species were dedicated to him, e.g. Octopus fontanianus d'Orbigny, 1834.
Fréminville, christophe Paulin de la Poix de (1787-1848). Known as Chevalier de Fréminville, he devoted his life to the French Navy. Performing archeology and natural history surveys, he served on the frigate Néréide off Western Africa. In mid-1822, the Néréide sailed to the Caribbean, visiting Martinique, Guadeloupe, and along the coast of the Islands of the Saints ("îles des Saintes"). In 1832 he described Eryon caribensis, a curious crab from Martinique that is actually a synonym of Symethis variolosa (Fabricius, 1793), and in 1835 he published a paper on the land crabs from the French Antilles that he had collected himself and given to the MNHN (Herpin 1913;Low et al. 2013). the managing committee of the "Muséum d'histoire naturelle de Genève", and its collections of Hymenoptera and Orthoptera became one of the best in the world (Hauser 1972). The CAA mentions exchanges between Saussure (MHNG) and the MNHN in 1858. (1811-1852). A French zoologist, malacologist and naval surgeon. He participated as naturalist and assistant surgeon on the voyage of La Bonite, which circumnavigated the globe (1836-1837) under the command of Auguste Nicolas Vaillant. After the death of Joseph Fortuné Théodore Eydoux (see under this name) in 1841, Souleyet spent ten years in Paris in order to himself complete the famous Eydoux et Souleyet's Voyage autour du monde […] de la Bonite. Souleyet died of yellow fever in Martinique (Laissus 1981;Bauchot et al. 1990Bauchot et al. , 1997.

souleyet, louis François auguste
Verreaux, Jules Pierre (1807-1873). French botanist and ornithologist and also a professional collector and trader in natural history specimens. He worked for the family business, Maison Verreaux, established by his father, Jacques Philippe Verreaux, at the Place des Vosges in Paris. It was the earliest known company specialising in natural history objects. The company, which funded numerous collection expeditions to various parts of the world, sold many specimens to the Muséum. In 1818, at the age of twelve, J. P. Verreaux accompanied his uncle Pierre Antoine Delalande (see under this name) on an expedition to the Cape of Good Hope, from where he brought back more than 131 000 specimens after three years of efforts. From 1820 to 1825 he received a proper scientific education at the Muséum, then returned to South Africa from 1825 to 1837. From 1842 to 1847, he carried out a long mission to Australia and Tasmania as an assistant naturalist for the Muséum, where he constantly collected samples. In 1864 he took over from Florent Prévot as assistant naturalist at the Muséum (Bauchot et al. 1990(Bauchot et al. , 1997Molina 2002). The name of various members of the Vearreaux family appears in the CAA: in 1837, "Verreaux" for material from Mauritius and Cape of Good Hope; in 1847, for a "lot of Verreaux collection from Tasmania, Australia (insects)"; "C. Verreaux" in 1865 for material from Gabon and for material from the coast of Africa near the Cape "given by E. Verreaux". Philip barker (1793-1854). An English botanist coming from an aristocratic family. He studied languages, botany, and geology at Harrow and Oxford. He collected plants in Italy, Spain, and Portugal, and was the first person to collect in the Tetuan Mountains of Morocco. En route to Brazil, he made what was intended to be a brief visit to the Canary Islands, but ended up stopping for a considerable time, returning after his Brazil expedition. Accompanied by Sabin Berthelot (see under this name), Webb collected specimens in the Canary Islands between 1828 and 1830. The results can be seen in the nine volumes of L'Histoire Naturelle des Iles Canaries, which he co-authored with Berthelot (see under this name), with contributions from other scientists (e.g. G. A. Brullé for the Crustacea). The text of this masterpiece took 20 years to be completed (Bauchot et al. 1990(Bauchot et al. , 1997. (1864-1961). He was a Swiss naturalist. He studied natural sciences at Basel and Bern, after obtaining his doctorate in philosophy in 1890 from the "Muséum d'histoire naturelle de Genève" as assistant entomologist to Henri de Saussure (see under this name). In 1894, he went to Java, from where he sent many insect specimens to Geneva. After a brief return to Switzerland, still as a collaborator with Saussure, he returned to Java in 1901, then visited Ceylon (now Sri Lanka). From 1912 to 1916, he led a forest research station in Juazeiro, Bahia, Brazil, where he conducted botanical excursions (Hauser 1972).