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NEW SPECIES AND NEW RECORDS OF ENDEMIC FRESHWATER CRABS FROM THE ATLANTIC FOREST IN SOUTHERN BRAZIL (ANOMURA: AEGLIDAE) Georgina Bond-Buckup, Carlos G. Jara, Ludwig Buckup, Marcos Pe´rez-Losada, Alessandra A.P. Bueno, Keith A. Crandall, and Sandro Santos (GBB, LB) Departamento de Zoologia, Instituto Biocieˆncias, Universidade Federal do Rio Grande do Sul, Av.Bento Gonc¸alves, 9500, 90501-970, Porto Alegre, RS, Brazil; (CGJ) Instituto de Zoologia, Casilla 567, Universidad Austral de Chile, Valdivia, Chile; (MPL) CIBIO, Centro de Investigac¸a˜o em Biodiversidade e Recursos Gene´ticos, Universidade do Porto, Campus Agra´rio de Vaira˜o, 4485-661, Vaira˜o, Portugal; (AAPB) Universidade Federal de Lavras, Departamento de Biologia, Campus Universita´rio, 37200-000 Lavras, MG, Brazil; (KAC) Department of Biology, Brigham Young University, Provo UT 84602-5255, USA; (SS, correspondence, [email protected]) Departamento de Biologia, Universidade Federal de Santa Maria, 97105-900, Santa Maria, RS, Brazil ABSTRACT Two new species of freshwater anomurans, Aegla pomerana and Aegla muelleri (Decapoda: Anomura: Aeglidae), are described from the Itajaı´ River basin, the major basin of the Atlantic range, located in the northeastern part of the state of Santa Catarina, Brazil. The new taxa can be distinguished from their congeners based on both morphological and molecular evidence (the nuclear gene 28S, and the mitochondrial genes 12S, 16S, COI, and COII). Based on molecular data, A. pomerana has a phylogenetic relationship with A. leptodactyla, but morphologically these two species differ in several characters. Aegla muelleri is a member of the same clade as A. leptochela, but several morphological characters distinguish the two species. New records of occurrence of A. jarai and A. odebrechtii, which occur in the same hydrographic basin, are provided.

KEY WORDS: Aegla, Anomura, biogeography, Brazil, Itajaı´ basin, molecular systematics DOI: 10.1651/09-3186.1

(Bond-Buckup, 2003). The first species of this family described from southern Brazilian continental waters was Aegla odebrechtii Mu¨ller 1876, recorded in streams that descend the Serra do Mar and enter the Rio Itajaı´ (BondBuckup and Buckup, 1994). Presently, six additional species of Aegla are recorded from the state: A. franciscana Buckup and Rossi 1977, A. platensis Schmitt1942, A. rossiana Bond-Buckup and Buckup 1994, A. spinosa BondBuckup and Buckup 1994, A. jarai Bond-Buckup and Buckup 1994 and A. parva Bond-Buckup and Buckup 1994. These last two have already been recorded in the basin of the Rio Itajaı´ (Bond-Buckup, 2003; Boos et al., 2006). In a recent survey of the Aeglidae of South America, the Rio Itajaı´ Basin was investigated in a most detailed manner, and the results are presented here. Two new species are described, and the distributions of A. jarai and A. odebrechtii are reviewed and extended.

INTRODUCTION The Atlantic Forest is considered one of the regions with the highest biodiversity in the world, a hotspot area (Mittermeier et al., 2004). The forest originally extended along almost the entire coast of Brazil, from the states of Piauı´ to Rio Grande do Sul. It covers an area of 1,300,000 Km2, which corresponds to 15% of the territory of the country, of which only 7% remains preserved. Even today the rivers and lakes of this forest shelters rich aquatic ecosystems, many of them threatened by the destruction of the gallery forests and consequent siltation of the water sources, by water pollution, and by the construction of reservoirs without due care for the environment. This intricate mosaic of basins is formed by rivers of national and regional importance, such as the Rio Itajaı´ in the state of Santa Catarina. The basin of the Rio Itajaı´ is the largest basin of the Atlantic drainage in the state of Santa Catarina, covering approximately 15,500 km2, equivalent to 16.5% of the area of the state (Fig. 1). The region has 53 municipalities occupying its territory, of which 47 sit within the basin, where about a million inhabitants live. In the Itajaı´ Valley, it is still possible to find remnants of the Atlantic Forest and watercourses in a good state of conservation, as is the case for the Serra do Itajaı´ National Park, in the municipality of Indaial. Living in rivers, creeks, and lakes, Aeglidae are considered good indicators of environmental quality, being found only in localities with clean, well-oxygenated water

MATERIAL AND METHODS The specimens were collected in watercourses of the Rio Itajaı´-Ac¸u´ Basin, during several sampling campaigns and were deposited in the Collection of Crustaceans of the Department of Zoology, Institute of Biosciences, Federal University of Rio Grande do Sul, Porto Alegre, Brazil (collection code: UFRGS) and in Museu de Zoologia da Universidade de Sa˜o Paulo (MZUSP). The descriptions of the new species were based on examination of the type-series. Measurements were made according to the method of Bond-Buckup and Buckup (1994), with the following abbreviations: ‘‘m’’ 5 males, ‘‘f’’ 5 females, ‘‘f ov’’ 5 ovate females, ‘‘j’’ 5 juveniles, CL 5

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Fig. 1. Drainage basin of the Rio Itajaı´, in the state of Santa Catarina. 1 5 Municipality of Pomerode; 2 5 Municipality of Indaial; 3 5 Municipality of Blumenau. total cephalothorax length (between the tip of rostrum and the midpoint of the posterior margin of the carapace), PCW 5 precervical width (between the left and right epibranchial margins), FW 5 frontal width (between the tips of the spines of the anterolateral angles of the carapace), AL 5 areola length, AW 5 areola width, RL 5 rostrum length (between the tip of the rostrum and the midpoint of the orbital margin. Morphometric Analysis For the description of the new species, the following body dimensions were recorded for all animals of the type series, and also for the nonparatype animals: TCL, CL, PCW, FW, AL, AW and RL. The morphometric ratios TCL/RL, CL/RL, PCW/FW and AL/AW were calculated for the set of animals from each locality. Molecular Analysis The molecular data and phylogenetic analyses presented in this study were obtained from Pe´rez-Losada et al. (2009). However, at the time of that publication, the taxonomic status of two species described here was unknown. Examination of the lots collected in the Rio Itajaı´-Ac¸u´ hydrographic basin allowed us to verify two new species and to add new localities of occurrence, with the respective geographical coordinates, of two species of previously described aeglids.

SYSTEMATICS Aeglidae Dana, 1852 Aegla pomerana n. sp. Bond-Buckup and Buckup (Fig. 2) Type-material.—Holotype male, Brazil, state of Santa Catarina, Pomerode, Sı´tio Mundo Antigo tributary in the Rio Itajaı´-Ac¸u´ Basin, 26u469220S, 49u119260W; 276 m a.s.l., 18.viii.1999, G. Bond-Buckup and L. Buckup col. (MZUSP 20463). Paratypes: 2 m, 1 f with juveniles, 2 j, same data as holotype (UFRGS 2679P); 1 m, same data as holotype, 02.i.2001, F. Faraco col. (UFRGS 2993); 4 m, 1 f, Campo Alegre, Joinvile, i.2002 (UFRGS 3262). Diagnosis.—Antero-lateral spine of carapace extending beyond base of cornea; protogastric lobes present; extraorbital sinus absent, rostrum triangular, slightly ligulate, slightly recurved distally, lacking carena on distal third, outer proximal margin of movable finger of cheliped with lobe; palmar crest of cheliped subrectangular; anterior angle of ventral margin of epimeron 2 unarmed; inner margin of ventral surface of ischium of cheliped with a distal tubercle.

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Description.—Carapace moderately compressed dorsoventrally, area of gastric region more elevated, dorsal surface scabrous, covered with punctae. Front wide; LPC/LF ratio of holotype male 1.87. Rostrum triangular, slightly ligulate, elevated in median portion, slightly recurved distally, without carena on distal third. Sub-rostral process developed; in profile, rostrum with ventral portion deeper than dorsal. Rostral carena begins between protogastric lobes, with two parallel rows of scales that do not reach apex; excavated in median portion. Lateral margins of rostrum with small scales. Orbits wide, deep, without orbital spine. Orbital margin with small sparse scales. Extraorbital sinus lacking. Antero-lateral angle of carapace projecting anteriorly in a spine, which extends beyond base of cornea. Outer and inner margins of antero-lateral lobe with sparse scales. First hepatic lobe delimited anteriorly by a distinct fissure; lateral margin with scales; second and third hepatic lobes not delimited, with only a slight indication; lateral margins with scales. Epigastric prominences little pronounced, surface irregular, of indefinite shape, elongated toward base of first hepatic lobe, with sparse scales. Protogastric lobes moderately elevated, anterior margin marked by row of scales. Transverse dorsal line sinuous. Areola subquadrate to subrectangular. CA/LA ratio of holotype male 1.5. Epibranchial area triangular, with an apical tubercle followed by scales. Lateral margins of anterior and posterior branchial areas with subequal scales. Anterior angle of ventral margin of epimeron 2 unarmed, with only small scales; ventrolateral margin slightly convex; posterior angle of ventral margin obtuse, unarmed. Epimera of third to sixth segments projecting; on third and fourth the lateral projection ornamented with a small apical scale. Telson divided by longitudinal suture. Anterior extremity of third sternite truncate, projected between the coxae of the exopods of the third maxillipeds. Fourth thoracic sternite elevated in median region, without ornament, lateral margins slightly recurved. Chelipeds subequal, hand subrectangular. Smaller chela delicate in appearance, elongated, covered by scales. Larger cheliped with more globose appearance, palm slightly more inflated in posterolateral region, covered with corneal scales. Palmar crest subrectangular, with margin ornamented with scaliform tubercles, indicating lobes. Pre-dactylar lobe forming a small angle with anterior margin of propodus, ornamented with scales. Fingers slender, covered by setae, scales, and scaliform tubercles. Proximal outer margin of movable finger with distinct lobe tipped with scales. Prehensile margins of fingers with scaliform denticles on their entire length, and with pronounced fitted opposed lobular teeth. Dorsal surface of carpus scabrous, with scales; inner margin with four spines, with distalmost spine being most developed; these spines with sparse scales on the lateral margins; small spine between the distal spine and the inner antero-lateral angle of the carpus; inner antero-lateral angle sub-obtuse, with apical spine; anterodorsal margin with scales. Distal part of

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Fig. 2. Aegla pomerana, n. sp. a: male holotype (MZUSP 20463), dorsal view; b: precervical portion of carapace, lateral view; c: ischium of cheliped, ventral view; d: third and fourth sterna, ventral view; e: second pleomere epimeron.

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dorsal surface with depression parallel to distal margin. Carpal crest pronounced, more elevated in proximal region, with clumps of three to five scales on elevations of crest; outer ventral angle of carpus with scales; ventral surface with one conical spine and tufts of setae. Dorsal margin of merus of cheliped with one spine, remainder of margin with scaliform tubercles; antero-dorsal margin with scales. Lateral surfaces with sparse punctae. Inner ventral margin of merus with one distal spine followed by scaliform tubercles; outer ventral margin with pronounced distal tubercle followed by scales. Dorsal margin of ischium with one spine; inner margin of ventral surface with small distal spine, followed by several spines along margin. Dorsal margin of dactylus, propodus, and carpus of second, third, and fourth pereiopods with rows of setae, scales, and scaliform tubercles arranged in longitudinal series, lending the surface a pubescent appearance. Variations.—The rostrum is more excavated in the larger paratypes, and the larger chela is more globose. Morphometry.—Holotype male with LC 16.89 mm. Paratypes (n 5 5) with mean LC 13.60 mm. A small-sized species. LPC/LF ratios of paratypes (n 5 3) ranging from 1.50 to 1.90. CA/LA ratios of paratypes varying little, from 1.7 to 1.8. Distribution.—Brazil: State of Santa Catarina, Rio Itajaı´Ac¸u´ Basin. Remarks.—The Bayesian phylogenetic analysis of the molecular data showed that A. pomerana belongs to Clade C as defined in Pe´rez-Losada et al. (2004). With regard to the relationship to other species, this was considered a sister-species of A. leptodactyla (Fig. 3), which occurs in the Rio Uruguay basin on the high plateau of Rio Grande do Sul and Santa Catarina. Nevertheless, there are several morphological differences between these two species. A. pomerana does not bear an extra-orbital sinus, whereas A. leptodactyla possesses this character. The rostrum of A. pomerana is only slightly ligulate, recurved, lacks a carina on the distal third, and ends without a spine on the tip; whereas this structure in A. leptodactyla is of medium length, and its distal end terminates in a spine. A. pomerana has wide chelipeds; its sister species has narrow chelipeds. The relationship of A. pomerana with species found in the Rio Uruguay basin, which at present has no direct connection to the Rio Itajaı´ basin, can be attributed to a common geological origin of the two basins, which is related to the formation of the Serra Geral in southern Brazil (Ribeiro, 2006). The uplift of the Serra Geral separated the Uruguay and Itajaı´-Ac¸u´ rivers creating the opportunity for new species to arise through allopatric speciation. Morphologically, A. pomerana resembles A. marginata Bond-Buckup and Buckup, 1994, in some aspects, particularly in the slightly arched antero-lateral border of the carapace. However, A. pomerana has a subrectangular palmar crest, and A. marginata has no palmar crest. Biology.—Unknown. Etymology.—Named in honor of the inhabitants of Pomerode, the type-locality of the species.

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Aegla muelleri n. sp. Bond-Buckup and Buckup (Fig. 4) Type-material.—Holotype male, Brazil, state of Santa Catarina, Indaial, Serra do Itajaı´ National Park, Rio Itajaı´Ac¸u´ basin, 27u019240S, 49u099060W; ii.2006, H. Boos col. (MZUSP 20464). Paratypes: 1 m, 6 f, same data as holotype (UFRGS 4117P); 2 m, 2 f, same data as holotype, 11.v.2001, H. Boos and G. Bond-Buckup col. (UFRGS 2982); id., 2 m, 21.ix.2002 (UFRGS 3570); id., juvenile (UFRGS 3571); 1 m, Cascata do Rio Passa Quatro, Monte Castelo, Rio Itajaı´-Ac¸u´ basin, 26o289140S, 50u119590W; 24.x.2000, A.A.P. Bueno, G. Bond-Buckup, C. Jara and M. Pe´rezLosada col. (UFRGS 2979); id., ib., ib., 11 m , 11 f , 1 f ov, 5 j (UFRGS 2980). Material Examined.—Brazil, Santa Catarina, Espingarda Creek, Serra do Itajaı´ National Park, Rio Itajaı´-Ac¸u´ basin, 27u01924.10S, 49u0996.70W; 20.vi.2002, H. Boos and K. Schacht col., 4 dissected specimens, (UFRGS 3385). Diagnosis.—Antero-lateral spine of carapace reaching base of cornea; protogastric and epigastric lobes absent; extraorbital sinus absent, rostrum triangular, tapered, slightly recurved distally, carinate, proximal outer margin of movable finger of cheliped without lobe; palmar crest of cheliped small, subrectangular; anterior angle of ventral margin of epimeron 2 with small scale; inner margin of ventral surface of ischium of cheliped with distal scaliform tubercle. Description.—Carapace strongly compressed dorsoventrally, dorsal surface scabrous, covered by punctations and scales; anterior dorsal region delimiting base of rostrum by V-shaped depression. Front wide; LPC/LF ratio of holotype male 1.73. Rostrum long, triangular, tapering, slightly recurved distally, carinate, distal part of rostrum slightly recurved. Sub-rostral process well developed; in profile, ventral portion of rostrum deeper than dorsal portion. Rostral carina begins at height of orbits, with two juxtaposed rows of scales that reach to apex; carina elevated in middle portion and very low in distal third. Lateral margins of rostrum scabrous. Orbits wide and deep, orbital spine indicated by an incision. Orbital margin scabrous. Extra-orbital sinus absent. Antero-lateral angle of carapace projecting anteriorly in a spine, extending beyond base of cornea. Outer and inner margins of antero-lateral lobe with sparse scales. First hepatic lobe delimited anteriorly by small incision; lateral margin smooth; 2nd hepatic lobe with small indication, and 3rd not delimited; lateral margins scabrous. Epigastric prominences lacking, surface scabrous. Protogastric lobes lacking, without indication. Transverse dorsal line sinuous. Areole quadrate. CA/LA ratio of holotype male 1.34. Epibranchial area triangular, with an apical tubercle followed by scales. Lateral margins of anterior and posterior branchial area with subequal scales.

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Fig. 3. Maximum likelihood and Bayesian trees as modified from Pe´rez-Losada et al. (2009). Posterior probabilities (if . 0.5) are indicated for each clade. Aegla pomerana and Aegla muelleri are presented in bold.

Anterior angle of ventral margin of epimeron 2 unarmed, indicated by small scale; ventro-lateral margin slightly convex; posterior angle of ventral margin obtuse, unarmed. Epimera of third to sixth segments projected; on the third and fourth the lateral projection ornamented with a small apical scale. Telson divided by longitudinal suture.

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Anterior extremity of third sternite triangular, projecting between coxae of exopods of third maxillipeds. Fourth thoracic sternite slightly elevated in medial region, without ornament, lateral margins slightly recurved. Chelipeds subequal, hand sub-rectangular. Chelae with delicate appearance, covered by scales. Larger cheliped

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Fig. 4. Aegla muelleri, n. sp. a: male holotype (MZUSP 20464), dorsal view; b: precervical portion of carapace, lateral view; c: ischium of cheliped, ventral view; d: third and fourth sterna, ventral view; e: second pleomere epimeron.

slightly more robust. Palmar crest subrectangular, with margin ornamented with scaliform tubercles, indicating lobes. Pre-dactylar lobe forming small angle with anterior margin of propodus, scabrous. Fingers slender, covered by sparse scales and setae. Outer proximal margin of movable finger without lobe. Prehensile margin of fingers with

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scaliform denticles along its entire length, and with fitted opposed lobular teeth. Dorsal surface of carpus scabrous, with scales; inner margin with three spines, the distalmost spine being the most developed of them; these spines without scales on their lateral margins; inner anterolateral angle projecting in apical spine; anterodorsal margin with

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scales. Dorsal surface distally with small depression parallel to distal margin. Carpal crest distinct, more elevated in proximal region; medial and distal portions with scales clumped in groups of three to five on elevations of crest; outer ventral angle of carpus obtuse, unarmed; ventral surface with small conical spine. Dorsal margin of merus of cheliped with elevations tipped with scales, which decrease in size proximally, followed by long setae; anterodorsal margin with sparse scales. Lateral surfaces with punctae and scales. Inner ventral margin of merus with a distal spine followed by scaliform tubercles and scales; prominent distal tubercle followed by scales on outer ventral margin. Dorsal margin of ischium with conical tubercle and long setae; inner margin of ventral surface of ischium elevation tipped with scale. Dorsal margins of dactylus, propodus, and carpus of second, third, and fourth pereiopods with rows of setae and scales arranged in longitudinal rows. Dorsal margins of second, third, and fourth pereiopods covered with long setae, lending the surface a pubescent appearance. Variations.—In some paratypes, the protogastric lobes are delimited anteriorly by sparse scales, although these are lacking in other specimens. Proximal outer margin of movable finger, in some specimens, bears scales suggesting a lobe. Other paratypes do not possess fitted opposed lobular teeth on the prehensile margin of the fingers, but only scaliform denticles on their entire length. The lot UFRGS 2979 has more-developed tubercles on the pereiopods, and the palmar crest is slightly more distinct. This is a small-sized species, comparable to A. violacea Bond-Buckup and Buckup, 1994 and A. leptodactyla. Morphometry.—Holotype male with LC 12.20 mm. The ovigerous female measured 13.29 mm LC. The mean LC of 25 paratypes was 11.52 mm, ranging from 8.0 to 15.33 mm. LPC/LF ratio of paratypes (n 5 10) ranging from 1.71 to 1.91, i.e., characterizing a wide front. CA/LA ratio of paratypes (n 5 11) ranging from 1.14 to 1.32. Etymology.—Named in honor of Fritz Mu¨ller, the first scientist to describe an aeglid from Brazilian waters. Biology.—Unknown. Distribution.—Brazil: state of Santa Catarina, Rio Itajaı´Ac¸u´ Basin. Remarks.—The new species A. muelleri resembles A. plana Buckup and Rossi, 1977 in the dorsoventrally compressed cephalothorax and the absence of the epigastric and protogastric lobes. However it is distinguished from A. plana by the shape of the chelipeds, which are wider in A. plana; by the configuration of the spines of the carpus; and by the shape of the palmar crest, which is more developed in A. plana. This latter species also possesses an extraorbital spine, which is lacking in A. muelleri. The total length of the cephalothorax of the paratypes (n 5 7) reached only 13.43 mm, showing it to be a small-sized species. The species occurs in sympatry with A. jarai in the Ribeira˜o Garcia in the Parque das Nascentes, an environ-

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mental preservation area that is among the few localities that shelter species in Atlantic Forest. The Bayesian phylogenetic analysis of the molecular data showed that A. muelleri belongs to Clade E as defined in Pe´rez-Losada et al. (2004). Aegla muelleri has a sister relationship to A. leptochela (Fig. 3), with which it shares the delicate form of the chelipeds and the convexity of the carapace. However, the two species differ in relation to the antero-lateral spine of the carapace, which in A. muelleri reaches the base of the cornea, and in A. leptochela does not. In A. muelleri, the extra-orbital sinus and the lobe of the movable finger are lacking, and the palmar crest is subrectangular; whereas in A. leptochela the extra-orbital sinus as well as the lobe of the movable finger are present, and the palmar crest is lacking. Aegla jarai Bond-Buckup and Buckup, 1994 Diagnosis.—Antero-lateral spine of carapace extending beyond middle of cornea; protogastric lobes subtle; rostrum very long in adult males, styliform, carinate along its entire length; anterior angle of ventral margin of epimeron 2 unarmed; outer proximal margin of movable finger of cheliped with lobe tipped with tubercle; palmar crest of cheliped disc-shaped, strongly excavated; inner margin of ventral surface of ischium of cheliped with one conical distal spine and up to three tubercles; dorsal margin of merus of second pereiopod with spine, followed by tubercles; ventral margin of merus of second pereiopod with scaliform tubercles. Formerly Known Distribution.—Brazil: Central and southern part of state of Santa Catarina, and northern part of state of Rio Grande do Sul (Bond-Buckup, 2003). New Records.—Brazil: State of Santa Catarina: Rio Itajaı´ basin: Rio Itajaı´-Ac¸u´: Ribeira˜o Encoro, Indaial, 13.v.2000, H. Boos col., 2 m, 5 f (UFRGS 2948); Ribeira˜o Espingarda, Indaial, 06.x.2000, H. Boos col., 2 m, 2 f (UFRGS 2991); Ribeira˜o Caete´, Cascata Ferdinand Schaback, Blumenau, 03.iii.2001, H. Boos col., 2 f, 1 j (UFRGS 3203). The occurrence of A. jarai in the sub-basin of the Rio Itajaı´-Ac¸u broadens its known geographical distribution to northeastern Santa Catarina. Biology.—Aspects of the growth of this species were characterized by Boos et al. (2006). Remarks.—The phylogeny of the aeglids shows that A. jarai belongs to Clade C, with an estimated divergence time of 33.2 6 1.8 MYA Pe´rez-Losada et al. (2004). The several populations of A. jarai form a nonmonophyletic group, according to these authors, indicating that new systematic studies on this species are needed. Aegla odebrechtii Mu¨ller, 1876 Diagnosis.—Antero-lateral spine of carapace reaching middle of cornea; protogastric lobes obsolete; rostrum of medium length in adult males, deflected and slightly recurved distally, without carina on distal third; anterior

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angle of ventral margin of epimeron 2 unarmed; proximal outer margin of movable finger of cheliped with lobe tipped with tubercles; palmar crest of cheliped subdisciform, excavated; inner margin of ventral surface of ischium of cheliped with one distal conical spine, one proximal smaller spine, and up to three tubercles between them. Formerly Known Distribution.—Brazil: The majority of records refer to the central region of the state of Santa Catarina, and the northern part of Rio Grande do Sul. In the Rio Itajaı´ basin, there is only the record of the holotype described from the Rio Itajaı´-Ac¸u, municipality of Blumenau. New Records.—Brazil: Santa Catarina: Rio Itajaı´ basin, a tributary of the Rio Itajaı´ do Sul (Fig. 1), Atalanta, 16.viii.1999, G. Bond-Buckup and L. Buckup col., 7 m, 2 f, 3 j (UFRGS 2676); Ribeira˜o Belchior, a tributary of Rio Itajaı´-Ac¸u, Gaspar, 14.vi. 1986, A da Silva col., 3 m, 2 f (UFRGS 2390). Biology.—Unknown. Remarks.—The phylogeny of the group, presented by Pe´rez-Losada et al. (2004), showed that A. odebrechtii belongs to clade C, with A. jarai as the sister species, and a divergence time estimated at 33.2 6 1.8 MYA.

ACKNOWLEDGEMENTS We thank our fellow biologists who participated in the sampling campaigns, especially Harry Boos Jr., Karin Schacht, and Fa´bio Faraco. We also thank CNPq for a productivity grant to GBB and SS, the US National Science Foundation (OISE-0530267, DEB-0075600) and the National Geographic Society for partial funding of this work.

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REFERENCES Bond-Buckup, G. 2003. Famı´lia Aeglidae, pp. 21-116. In, G.A.S. Melo (ed.), Manual de identificac¸a˜o dos Crustacea Decapoda, de a´gua doce do Brasil. Editora Loyola, Sa˜o Paulo. ———, and L. Buckup. 1994. A famı´lia Aeglidae (Crustacea, Decapoda, Anomura). Arquivos de Zoologia 32: 159-347. Buckup, L., and A. Rossi. 1977. O geˆnero Aegla no Rio Grande do Sul, Brasil (Crustacea, Decapoda, Anomura, Aeglidae). Revista Brasiliero de Biologia 37: 879-892. Boos Jr., H., D. Silva-Castiglioni, K. Schacht, L. Buckup, and G. BondBuckup. 2006. Crescimento de Aegla jarai Bond-Buckup and Buckup (Crustacea, Anomura, Aeglidae). Revista Brasileira de Zoologia 23: 490-496. Mittermeier, R.A., P.R. Gil, M. Hoffmann, J. Pilgrim, T. Brooks, C.G. Mittermeier, J. Lamoreux, and G.A.B. Fonseca. 2004. Hotspots Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial Ecoregions. CEMEX/Agrupacio´n Sierra Madre. 432 pp. Mu¨ller, F. 1876. Aegla odebrechtii n. sp. Jenaische Zeitschrift fu¨r Naturwissenschaft, n.s., Jena 10(3): 13-24. Pe´rez-Losada, M., G. Bond-Buckup, C.G. Jara, and K.A. Crandall. 2004. Molecular systematics and biogeography of the southern South American fresh-water ‘‘crabs’’ Aegla (Decapoda: Anomura: Aeglidae) using multiple heuristic tree search approaches. Systematic Biology 53: 767-780. ———, ———, ———, and ———. 2009. Conservation Assessment of Southern South American Freshwater Ecoregions on the Basis of the Distribution and Genetic Diversity of Crabs from the Genus Aegla. Conservation Biology, 23: 692-702. Ribeiro, A. C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotropical Ichthyology 4: 225-246. Schmitt, W.L. 1942. The species of Aegla, endemic South American crustaceans. Proceedings of the United States National Museum 91: 431-524.

RECEIVED: 9 June 2009. ACCEPTED: 18 November 2009.