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 (ecampos@uabc.edu.mx)
Corresponding editor: Ingo Wehrtmann

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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.

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INTRODUCTION

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.

MATERIALS AND METHODS

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.

 

RESULTS

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)

Diagnosis

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

Synonymy

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.

Color

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

Measurements

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.

DISCUSSION

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.
Pinnotherespisum (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
chilensis (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).

 

ACKNOWLEDGMENTS

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.

 

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Received: 20 April 2016;
Accepted: 13 January 2017