Research Article |
Corresponding author: Cristiano Sampaio Costa ( csampaioc@gmail.com ) Academic editor: Ângelo Pinto
© 2018 Cristiano Sampaio Costa, Amazonas Chagas-Junior, Ricardo Pinto-da-Rocha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Costa CS, Chagas-Jr A, Pinto-da-Rocha R (2018) Redescription of Epiperipatus edwardsii, and descriptions of five new species of Epiperipatus from Brazil (Onychophora: Peripatidae). Zoologia 35: 1-15. https://doi.org/10.3897/zoologia.35.e23366
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Epiperipatus edwardsii (Blanchard, 1847) is redescribed based on fresh material collected near the type locality. Additionally, five new species of velvet worms of the genus Epiperipatus Clark, 1913 from Brazil are described: Epiperipatus hyperbolicus sp. nov., Epiperipatus lucerna sp. nov., Epiperipatus titanicus sp. nov. (Holotypes deposited in MNRJ: Murici, Alagoas State), Epiperipatus beckeri sp. nov. (Holotype female deposited in MNRJ: Camacan, Bahia State) and Epiperipatus marajoara sp. nov. (Holotype male deposited in MZUSP: Breves, Marajó island, Pará State). The peculiar shape of the primary papillae (artichoke-like) of E. titanicus sp. nov. and E. beckeri sp. nov. is documented for the first time. Epiperipatus titanicus sp. nov. has the largest number of pairs of oncopods (38 for males and 39 for females) among the Brazilian Onychophora. We also provide an identification key for Brazilian species of Epiperipatus.
French Guyana, neotropics, onychophorans, Peripatus , taxonomy.
Epiperipatus Clark, 1913 is the most diverse genus of Neotropical Peripatidae. Currently, 26 species are known in Central and South America (
Neopatida is the largest group of Peripatidae from the Neotropics. This taxonomically obscure group comprises the widespread genus Macroperipatus Clark, 1913, type species Peripatus torquatus von Kennel, 1883, from Trinidad, and Epiperipatus, type species Peripatus edwardsii Blanchard, 1847, from French Guiana. Those genera were included in a phylogenetic analyses of
Epiperipatus edwardsii is widespread in the Neotropics, distributed on a large area from Panama, Trininad & Tobago, Venezuela, French Guiana, Suriname, Colombia and Brazil (
Based on findings of
In total, we examined 65 specimens collected between 2008 and 2014. The material examined is deposited in the following institutions: Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP); Universidade Federal de Minas Gerais, Belo Horizonte, Brazil (UFMG); Natural History Museum, London, United Kingdom (NHM); Collection of the Evolution et Diversité Biologique laboratory at Université Toulouse III Paul Sabatier, Toulouse, France (GF). Live specimens were photographed (mainly for color and body patterns) with SONY Cybershot DSC-HX1 with built-in flash, or Canon EOS Rebel XS with macrolens and flash circular cameras. The color of live specimens was described following the standard names of the 267 Color Centroids of NBS/ISCC Color System (
We obtained the stacked images using a Leica MZ125 Stereomicroscope with built-in Leica DFC 290 camera, Leica Light source CLS100X series 00373 and assembled through of software Leica Application Suite version 3.3.0. We compiled descriptions, and dissection of dead specimens using a stereomicroscope. The morphological descriptive nomenclature follows the terminologies of
Dorsal integument, jaws, legs and some antennae were photographed using Scanning Electron Microscopy (SEM). All tissues were cleaned, subjected to critical point and sputter coating. In the cleaning process, the soft parts were washed in a solution of hypochlorite 3% diluted in distilled water. The hard parts (for example jaws) were cleaned in distilled water and Ethylenediaminetetraacetic acid (or EDTA) heat solution in sonicator processor. For critical point, the tissues were dehydrated using an increasing ethanol series: 30%, 50%, 70%, 80%, 90% and 100%.
These tissues were critical point dried and mounted in SEM stubs with biadhesive carbon tape, in the critical point machines Baltec CPD 030. The sputter coating was completed using the machines Denton Desk IV gold sputter model and Balzer SCD 50 sputter coater. Generally, on average, 5 to 10 nm of gold/paladium or platinum/palladium were deposited over the tissues. We took SEM images using microscopes JEOL JSM-6390LV and DSM 940 Zeiss scanning electron. The Backscatter electron detector (BSD) function, when available on the microscope, was used to avoid the charging effects on the tissues.
The descriptions of new species are based on type series instead of just in the holotype (compound description). A full description is only provided for the first described species, E. titanicus, subsequent descriptions are comparative and complementary to that of E. titanicus.
Peripatus (Epiperipatus)
Clark, 1913: 17;
Epiperipatus
Clark, 1937: 2;
Peripatus edwardsii Blanchard, 1847 by original designation.
Dorsal papillae with roundish insertion. Bases of primary papillae with four to eighteen scale ranks. Apical pieces are conical, cylindrical, or rarely flat. Usually four complete and sometimes, rarely vestiges of a fifth spinous pad on the fourth and fifth legs. Nephridial tubercle on fourth and fifth pairs of legs, between third and fourth spinous pads. One to tree pregenital legs with crural papillae (male only).
Peripatus edwardsii Blanchard, 1847: 140.
Peripatus (Epiperipatus) edwardsii
:
Epiperipatus
edwardsii
:
The background color of the body of this species is in the purple spectrum; dorsal papillae round, primary papillae bearing asymmetrical apical piece, four complete spinous pads on the fourth and fifth legs (the last broken in some specimens).
Measurements (mm): Type: length 35.0; width 4.0; height 3.0 (based on
Epiperipatus edwardsii,GF180312HC003-06: (1) dorsal papillae arrangement; (2) dorsal view of the distal half of the body depicting diamond shaped pattern over the dorsomedian furrow; (3) foot in ventral view; (4) primary papillae in posterior view depicting differences in the number and shape of the scales (B and P); (5) outer blade in first plan of right jaw; (6) right fourth leg in ventral view depicting spinous pad undivided (I) and divided (IV) into two unqueal parts. (AT) Accessory tooth, (B) basal piece, (DR) series of denticles, (I) first spinous pads, (IJ) inner blade, (IV) last spinous pads, (nt) nephridial tubercle, (OJ) outer blade, (P) apical piece, (PP) prolateral foot papillae, (RP) retrolateral foot papillae. Scale bars: 1, 5 = 100 µm, 2 = 5 mm, 3 = 30 µm, 4 = 20 µm, 6 = 200 µm.
Head. No evident structures or color patterns on head. Antennae with 44 rings: antennal tip composed of seven broad rings, excluding the terminal disc on top. Antennal body with alternating sequence of narrow and broad rings at least up to the eighteenth. Eyes and frontal organs present on ventrolateral region of antennal base. Frontal organs as long as four fused antennal papillae. Mouth opening surrounded by small, anterior, unique lobe, and seven flanked lobes decreasing in size from anterior to posterior ends of the mouth. Dental formula of inner and outer jaws, respectively: 1/1 and 1/1/10, with second accessory tooth of inner jaw vestigial (Fig.
Legs. Male with 30 pairs of legs. Ventrally, fourth and fifth pairs of legs with four complete spinous pads (the fourth are sometimes broken) without vestige of fifth spinous pad (Fig.
Sexual dimorphism. Male pregenital leg without crural papillae (these organs had not been previously described for this species). Anal glands inconspicuous (male); represented by two pores on ventral portion of anal aperture.
Material examined. 1♂, French Guiana, Cayenne, Nouragues Field Station, no further data (GF180312HC003-06). The holotype, deposited at the Muséum National d’Histoire Naturelle, Paris, France (MNHN), was not examined because the curator did not reply to our loan requests.
Distribution. French Guyana: Cayenne and adjacency of Nouragues Field Station (Bélizon, Bita, Coco).
Remarks. The type of Epiperipatus edwardsii, deposited at the Muséum National d’Histoire Naturelle, Paris, France (MNHN) was not examined. We tried to contact the curator to arrange a loan of this material, but the curator did not reply. Epiperipatus edwardsii is the most frequently misidentified species of Epiperipatus and for this reason some records of its distribution (e.g.,
According to our observations, the diamond patterns are a general feature in most species of Epiperipatus. Morphologically, the diagnostic features to delimit E. edwardsii are (1) dorsal papillae over the plicae and never on the folds, and (2) primary papillae usually separated by one or two accessory papillae (however in some cases they are close to each other). Also, E. edwardsii is supported by molecular data as demonstrated in Maximum Parsimony and Maximum Likelihood analyses (e.g.,
The other records of E. edwardsii found in the literature, for instance from Panama, Venezuela, Colombia and Brazil (
Epiperipatus
sp. 1:
Epiperipatus species with symmetrical, reduced or hidden apical piece. Body large, 29 up to 76 mm and 36 to 39 pairs of legs.
Measurements. Males: length 29–41; width 1.0–3.0; height 1.0–2.5. Females: length 16–76; width 1.0–4.0; height 1.0–6.0. Background color of body in vivo is dark grayish red (20), with dorsal wavy band of light grayish red (18) and overlaid by diamond-shape marks, the same color as background color of body (Fig.
Shape of primary papillae (holotype, except when mentioned) of Epiperipatus beckeri sp. nov., E. piperipatus lucerna sp. nov., Epiperipatus marajoara sp. nov., Epiperipatus hyperbolicus sp. nov. and Epiperipatus titanicus sp. nov. (7) Epiperipatus titanicus sp. nov. (MNRJ 0057); (8) Epiperipatus marajoara sp. nov. (MZUSP 0022); (9) Epiperipatus lucerna sp. nov. MNRJ 0101; (10) Epiperipatus beckeri sp. nov. (MNRJ 0045); (11–12), Epiperipatus hyperbolicus sp. nov. (MNRJ 0104). Scales bars: 7, 12 (also in the small box of Fig.
Epiperipatus titanicus sp. nov. habitus, dorsal plicae, spinous pad and jaws: (13) Living specimen in dorsal habitus, paratype; (14) dorsal papillae arrangement of the holotype; (15) paratype blades; (16) holotype blades with the accessory tooth broken in outer jaws; (17) spinous pad and nephridial tubercles of paratype. The Roman numbers indicate the first and last spinous pad of the leg. Black arrows point vestiges of a fifth spinous pad; (15–16) jaws with outer blade in first plane in the both figures. Scales bars: 13 = 2.7 mm, 17 = 200 µm, 14– 15 = 100 µm, 16 = 50 µm.
Head. Holotype and paratype with 44–45 antennal rings, respectively. In both specimens, antennal tip composed of seven broad rings (excluding the terminal disc on top) and followed by alternating narrow and broad rings until the eighteenth ring. Mouth opening surrounded by anterior oral lobe (unpaired lip) and seven pairs of oral lips. Jaws with two blades composed of one long curved main tooth followed by one accessory tooth (Figs
Two or three pregenital legs with one crural papilla (male) each. Anal glands inconspicuous (male); represented by two pores on anterior margin of anal aperture.
Holotype. ♀, BRAZIL, Alagoas State: Murici, Estação Ecológica de Murici (Mata da Bananeira area), Lo-Man-Hung, N.F. leg. (MNRJ 0057). Paratypes. ♂, idem, 11–18.xiii.2009, Costa, C.S., Giupponi, A. and Chagas-Jr leg. (MNRJ 0035); 2 ♀, same date but 21.vii.1995, Freitas, L.M. and Natali, M.S. leg. (DZUFMG-ONY 0008 and 0009); 9♀, 4 ♂ and 1 unsexed, same data but, 11–18.xii.2009, Costa, C.S., Guipponi, A. and Chagas-Jr, A, leg (MNRJ 0053); 1♀, same data but, 26–29.vii.2012, Costa, C.S., Alcântara, D.M.C., Dias, P.S. and Nihei,S.S. leg (MZUSP 0010).
Distribution (Fig.
Etymology. The specific name titanicus, titanica, titanicum (L., “titanic”, adjective) refers to the large size of the females, and the great number of pairs of oncopods, more numerous than in any other Brazilian species of onychophorans.
Remarks. The number of leg pairs in this species varies from 36 to 39, and these numbers overlap in males and females, 36 to 38 in males and 36 to 39 in females.
Apical piece cylindrical, asymmetric, have of two scale ranks on back side and three on front side; 29 to 30 pairs of legs.
Measurements. Male: length 13–29; width 1.0–3.0; height 1.0–1.5. Female: length 23–44; width 1.0–3.0; height 2.0–4.0. Color (living specimens). In vivo background color of body on dorsal portion, arrangement of pale primary papillae, and ventral region as described for E. titanicus. Description of body. Dorsal papillae aligned on top of folds, with two primary papillae close together or separated by one to five accessory papillae. Dorsal papillae with conical basal piece, composed of overlapping lanceolate scales. Primary papillae regular in size, at their bases four to eight scale ranks (Fig.
Epiperipatus lucerna sp. nov. and Epiperipatus hyperbolicus sp. nov.. Epiperipatus lucerna sp. nov., dorsal plicae, spinous pad and jaws: (18) dorsal papillae arrangement over the plicae (holotype); (19) primary papillae, holotype. Note the scales ranks on the basal piece; (20) spinous pad and nephridial tubercles of paratype. Epiperipatus hyperbolicus sp. nov., dorsal plicae, spinous pad and jaw, paratype images: (21) dorsal plicae on top view. Incomplete plicae position indicate by the arrow head; (22) spinous pad and nephridial tubercles. The Roman numbers indicate the first and last spinous pad of the leg; (23) jaw. The inner and outer blades are illustrated in the large and small boxes. Scales bars: 18, 21 = 500 µm, 19 = 20 µm, 20 = 200 µm, 22 = 100 µm, 23 (also in the small box) = 50 µm.
One or two pregenital legs with crural papillae (male) with one crural papilla each (structure not identifiable in some adult males examined). Anal glands inconspicuous (male); represented by two pores on anterior board of anal aperture.
Holotype. MNRJ 0101, ♂, Brazil, Alagoas state, Murici, Estação Ecológica de Murici (Mata da Bananeira area), 11–18.xii.2009, Costa, C.S., Giupponi, A. and Chagas–Jr leg. Paratypes. MNRJ 0102, ♀, idem; MNRJ 0103, 13♀, 6 ♂ and 3 unsexed, Brazil, Alagoas state, Murici, Estação Ecológica de Murici (Mata da Bananeira area), 11–18.xii.2009, Costa, C.S., Giupponi, A. and Chagas–Jr leg; MZUSP 0011, 1♀, idem, 26–29.vii.2012, Costa, C.S., Alcântara, D.M.C., Dias, P.S. and Nihei,S.S. leg.
(Fig.
The specific name lucerna (L., “lamp”, feminine noun in apposition) refers to the candle shape of the apical piece of the primary papillae of this species.
Dorsal papillae with conical and small basal piece composed of truncated robust scales and primary papillae of regular size with six or seven scale ranks at base and robust and asymmetric spherical apical piece. Legs with 23 to 25 pairs.
Measurements (based on a single male): length 26, width 2.0 and height 3.0. Females: length 22–33, width 17–2.0 and height 2.0–3.0. Color (living specimens). Background color of dorsal body, arrangement of pale primary papillae, and ventral region as in Epiperipatus titanicus sp. nov. Description of body. Dorsal papillae aligned on top of folds, two primary papillae occurring close together or separated by one to three accessory papillae. Dorsal papillae with conical basal piece, composed of truncated scales (Fig.
Head. In holotype antennae with 27 rings (28 rings in paratype, some females with 36, 37 and 41 antennal rings). In both holotype and paratype the antennal tips composed of seven broad rings and terminal bud. Antennal tip followed by alternating narrow and broad rings until sixteenth ring. Jaw blades as in Epiperipatus titanicus sp. nov. with dental formula of holotype and paratype as follows: 1/1/7 and 1/1/11 (Fig.
Epiperipatus beckeri sp. nov. and Epiperipatus marajoara sp. nov. Epiperipatus beckeri sp. nov., dorsal body background and papillae and jaw, holotype, MNRJ 0045: (24) body background of dorsal side; (25) dorsal plicae and papillae arrangement. Note the incomplete plicae on the right side of the picture; (26) accessory papilla in posterior view. Note slight different shape between the accessory and primary papillae (see the Fig.
One or two pairs of pregenital legs present (male) with one crural papilla on each. Anal glands inconspicuous (male); represented only by two pores on anterior margin of anal aperture.
Holotype. MNRJ 0104, ♂, Brazil, Alagoas State, Murici, Estação Ecológica de Murici (Mata da Bananeira), 11–18.xii.2009, Costa, C.S., Giupponi, A. and Chagas-Jr leg. Paratypes. MNRJ 0105, ♀, idem; MNRJ 0106, 2♀, Brazil, Alagoas State, Murici, Estação Ecológica de Murici (Mata da Bananeira), 11–18.xii.2009, Costa, C.S., Giupponi, A. and Chagas-Jr leg.
(Fig.
The specific name hyperbolicus, hyperbolica, hyperbolicum (Latinized adjective “exaggerated”, from Greek ὑπερβολή, plus suffix -icus) refers to the large spherical shape of the apical piece of the primary papillae of this species.
Dorsal background color of body light orange, with dark orange dorsomedian furrow, and antennae purple brownish. Apical piece of primary papillae symmetrical, reduced, not hidden basally.
Compound description. Measurements: length 35–53, width 2.0–4.0, height 4.0. Holotype: length 53, width 4.0, height 2.0. Color (living specimens). Background color of body of the specimen is moderate yellowish pink (29) with a mid-longitudinal pale orange yellow (73) wavy stripe interrupted by well-defined background color cardioid patches (Fig.
Legs. Twenty-eight to thirty (juveniles included). Holotype with 30 pairs of legs. Nephridial tubercles between third and fourth spinous pads, connected by strait isthmus on top with third spinous pad. Two prolateral and one retrolateral foot papillae on feet of fourth and fifth legs.
Epiperipatus marajoara sp. nov., Epiperipatus cratensis, E. brasiliensis and E. paurognostus. Epiperipatus marajoara sp. nov., MZUSP 0022, holotype: (31) spinous pad and the nephridial tubercle of the fifth leg; (32) inner and outer blade of right jaw. The accessory tooth is broken in the outer blade. (33, 35) Epiperipatus cratensis, MZUSP 0083, dorsal papillae arrangement with dorsomedian furrow on the top of image and primary papillae in lateral view, respectively. (34) E. brasiliensis, MZUSP 0121, primary papillae in lateral view. (36) E. paurognostus, UFMG 0184, dorsal papillae arrangement. Note all dorsal papillae are proeminent. Scale bars 31, 32 = 30 µm, 33 = 500 µm, 34 = 20 µm, 35 = 100 µm, 36 = 40 µm.
Males unknown.
Holotype. ♀, Brazil, Bahia State, Camacan, Reserva Particular do Patrimônio Natural Serra Bonita, 11–15.vi.2009, Chagas-Jr, A., Dill, V., Giupponi, A., Pedroso, D. & Kury, A. Leg (MNRJ 0045),. Paratypes. 1 ♀ and 4 juveniles, Brazil, Bahia State, Camacan, Reserva Particular do Patrimônio Natural da Serra Bonita, 14.ii to 12.iii.2009, M. Teixeira Junior leg. (MZUSP 0017).
(Fig.
The specific name beckeri (German surname, noun with Latin ending -i of the genitive case) honoring the naturalist and entomologist Vitor O. Becker, who created and is responsible for the administration of Reserva Particular da Serra Bonita, Camacan, Bahia State.
The main differences between juveniles and adults are the color and the kind of dorsal papillae they display. A juvenile was photographed in vivo with background color of body very dark purplish red (260), without diamond areas. The other post-fixed juvenile darker than adults, with background color of body dark brown (55), diamond areas brownish orange (54) and legs on dorsal region pale yellow (89).
The regular primary papillae and accessory papillae have similar size in the adult. In the juveniles, the largest pale primary papillae are more diverse on the dorsal plicae. These primary papillae have a spherical everted and robust apical piece, their arrangement forming longitudinal lines similar to the condition found in the holotype.
Peripatus
simoni
: Bouvier, 1900: 753; 1905: 315;
Epiperipatus
simoni
:
This species has dorsal background color of body in the brown spectrum, dorsal diamond pattern inconspicuous or absent, roundish dorsal papillae, primary papillae with well-developed base, and a cylindrical apical piece.
Compound description. Measurements. Females. Length 40, 50 and 65 (specimens from NHM examined by Bouvier in his monograph). Males. Length 27–34, width 2.0–7.0, height 1.0–6.0. The single female is: length 33, width 2.8, height 1.2. Color (living specimens). Background color of body Light Grayish Brown (60) overlaid with brownish pink (33) diamond-shaped makers. Dorsalmedian furrow moderate brown (58), and antennae grayish reddish brown (46) (Figs
Head. Antennae composed of 43 to 47 rings. Antennal tip composed of seven broad rings, except the terminal disc on top, followed by sequence of narrow and broad rings until at least the eighteenth antennal ring. Jaws with two blades and both composed of one long curved main tooth followed by one accessory tooth. Dental formula of outer and inner for holotype and paratype are: 1/1 and 1/1/8 (Fig.
Sexual dimorphism. One or two pregenital legs with one or two crural papillae (male), absent in females. Anal glands inconspicuous (male); represented only by two pores on anterior margin of anal aperture, absent in females.
Holotype. ♂, Brazil, Pará State, Island of Marajó, Breves, Reserva de Extrativista Mapuá, Breves; 4–8.xii.2012; Cristiano Sampaio Costa leg. (MZUSP 0022). Paratypes. Same data as holotype. ♀ (MZUSP 0027). 4 ♂ (MZUSP 0023-0026). 3♀, Breves, Marajó Island, Brazil, Amazon, 96.5.14.6-8 (NHM # 1004943, 1004944, 1004945).
(Fig.
The specific name marajoara (demonymic noun in apposition; invariable), honors the marajoara people, natives from the Island of Marajó.
This species was discovered comparing the descriptions with bona fide specimens of E. simoni from NHM and specimens collected from the Island of Marajó, Pará State, Brazil. The type locality of E. simoni is Caracas in Venezuela, with records in Breves, Island of Marajó (Pará State) and Rio de Janeiro State (see
Since 2010, the number of species of Epiperipatus has increased through descriptions of new species and new combinations. After the five new species erected in this work, the genus became the most diverse of Neotropical onychophorans, now including 31 species. In Brazil, Epiperipatus is also the most diverse genus of Peripatidae, with 14 species (excluding E. edwardsii and E. simoni, Fig.
As stated by
Species | Morphology of the apical piece | Distribution of ranks of scales1 | Range of scale ranks2 | Incomplete plicae | Number of oncopods |
E. acacioi | Spherical | Asymetrical | 2 or 3 | Present | 24–26 (♂); 26–29 (♀) |
E. adenocryptus | Conical | Asymetrical | 2 or 3 | Present | 26 and 27 (♂); 28–30 (♀) |
E. brasiliensis | Conical | Symetrical | 2 | Absent | 29 (♂); 31 and 32 (♀) |
E. cratensis | Spherical | Asymetrical | 2 or 3 | Absent | 30–33 (♂); 33–34 (♀) |
E. diadenoproctus | Conical | Asymetrical | 2 or 3 | Present | 26–28 (♂); 29 and 30 (♀) |
E. machadoi | Conical | Asymetrical | 2 or 3 | Present | 27–29 (♂); 28–31 (♀) |
E. ohausi | Spherical | Asymetrical | 3 | Present | 26–28 (♂); 27–29 (♀) |
E. paurognostus | Conical | Asymetrical | 2 or 3 | Present | 26 and 27 (♂); 27–29 (♀) |
E. tucupi | Conical | – | – | Absent | 34 and 35 (♀) |
E. beckeri sp. nov. | Conical reduced | Symetrical | 1 | Present | 28–30 (♀) |
E. hyperbolicus sp. nov | Spherical robust | Asymetrical | 3 | Present | 23 (♂); 24 and 25 (♀) |
E. lucerna sp. nov. | Conical | Asymetrical | 2 | Present | 27–29 (♂); 29–31 (♀) |
E. marajoara sp. nov. | Conical | Asymetrical | 2 | Present | 27 (♂); 31 (♀) |
E. titanicus sp. nov. | Conical reduced | Symetrical | 1 | Present | 36 and 38 (♂); 36–39 (♀) |
However, the five new erected species are herein assigned to Epiperipatus because they share, with other species in the genus, the rounded insertion of the dorsal papillae, and distinct nephridial tubercles between the third and fourth spinous pads. They are also nested within of Epiperipatus based on molecular synapomorphyes of found in COI, 12S rRNA, 16S rRNA and 18S rRNA sequences (
Our study shows new characters to the taxonomy of Peripatidae. For instance, the few scales ranks on apical piece (see E. titanicus and E. beckeri), and the larger shape apical piece (see E. hyperbolicus). In resume, the new species differ from each other by the shape of the apical piece on the primary papillae. The apical piece of the primary papillae in E. titanicus is smaller than in E. beckeri. As for E. lucerna sp. nov. and E. marajoara sp. nov., they have conical apical piece, but E. marajoara sp. nov. has a smaller number of pairs of oncopods (27–31 pairs) and inconspicuous diamond pattern on the dorsal portion of the body. Among the five new species, E. hyperbolicus sp. nov. is distinguished by the particular shape of the apical piece of the primary papillae. Only in this species a robust and asymmetric spherical apical piece was observed.
Additionally, E. lucerna sp. nov., E. hyperbolicus sp. nov. and E. titanicus sp. nov. are the first species described in Brazil from the same locality, Estação Ecológica de Murici (Mata da Bananeira) in the Alagoas State, Brazil. This region is highland humid forest inserted in the realm of the Caatinga, and is really important to the understanding of the distribution and biogeographical history of onychophorans in the Neotropical region (Vasconcellos et al. 2004).
Due to the conservative morphology of Neotropical onychophorans, many specimens previously identified as E. edwardsii in the literature are not actually E. edwardsii. The characters that have been most often misinterpreted are the shape and distribution of the dorsal papillae, the dorsal diamond pattern, and the number of oncopods. A recent phylogenetic study revealed that E. edwardsii occurs only in Cayenne, or near this locality (
1 | Complete folds (Fig. |
2 |
1’ | Incomplete folds (Fig. |
3 |
2 | Symmetrical conical apical piece (Fig. |
E. brasiliensis Bouvier, 1899 |
2’ | Asymmetrical conical apical piece (Fig. |
E. cratensis Brito et al., 2010 |
2” | Asymmetrical spherical well-developed apical piece (comparable with E. cratensis), 34 and 35 pairs of legs (females) | E. tucupi Froehlich, 1968 |
3 | Conical apical piece (Figs |
4 |
3’ | Spherical apical piece (Fig. |
5 |
4 | Conical normal-sized apical piece (Fig. |
6 |
4’ | Conical/cylindrical reduced (comparable with E. edwardsii) apical piece (Figs |
7 |
5 | Spherical regular apical piece (Fig. |
8 |
5’ | Spherical robust (comparable with E. edwardsii) apical piece (Fig. |
E. hyperbolicus sp. nov. |
6 | Asymmetrical apical piece with cylindrical shape (Fig. |
E. marajoara sp. nov. |
6’ | Asymmetrical apical piece with conical candle shape (Fig. |
E. lucerna sp. nov. |
7 | Symmetrical reduced or hidden apical piece (Fig. |
E. titanicus sp. nov. |
7’ | Symmetrical reduced apical piece, dorsal papillae aligned on top of folds (Fig. |
E. beckeri sp. nov. |
8 | Peripheral accessory papillae frequently occur over the flank (Fig. |
9 |
8’ | Peripheral accessory papillae rare over the flank ( |
10 |
9 | Background brownish, 27 to 29 pairs of legs and 28 to 31 (females) | E. machadoi (Oliveira & Wieloch, 2005) |
9’ | Background dark red, 24 to 26 pairs of legs (males) and 26–29 (females) | E. acacioi (Marcus & Marcus, 1955) |
10 | Conspicuous anal gland (only males, |
11 |
10’ | Unconspicuous anal gland (only males, |
12 |
11 | Male anal glands with two well-developed pores (like light spots) on the anterior board of anal aperture ( |
E. diadenoproctus Oliveira et al., 2011 |
11’ | Male unconspicuous anal gland, which was bean-shaped, hardly visible ( |
E. adenocryptus Oliveira et al., 2011 |
12 | Only primary papillae are prominent (Chagas-Jr & Sampaio 2014: fig. 6), primary papillae with robust apical piece (Chagas-Jr & Sampaio 2014: figs 8, 9), 26 to 28 pairs of legs (male) and 27 to 29 (female) | E. ohausi Bouvier, 1900 |
12’ | All dorsal papillae are prominent (Fig. |
E. paurognostus Oliveira et al., 2011 |
We are grateful to Jerome Murienne, who provided specimens of E. edwardsii for the present study. We also acknowledge partial support funding such as Labex CEBA (ANR-10-LABX-25-01) to collect E. edwardsii. This work was partially supported by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2011/20211-0, 2012/02969-6 and 2014/20557-2) to C.S.C. and R.P.R. We are grateful to Adriano B. Kury who provided support with the Latin grammatical forms. Beka Buckman, Brittany Damron and Denis Jacob provided help and comments to the manuscript. We acknowledge the support of the Center for Scanning Electron Microscopy of the Museu Nacional, Universidade Federal do Rio de Janeiro, the electron microscope laboratory at Instituto de Biociências of the Universidade São Paulo, the electron microscope laboratory at Museu de Zoologia da Universidade São Paulo, and the Center for Nanoscale Systems, Harvard University for the SEM images.