ZOOLOGIA 26 (3): 553561, September, 2009

doi: 10.1590/S1984-46702009005000004

Brazilian species of Onychophora with notes on their taxonomy and distribution

Cristiano Sampaio-Costa

1, 3

; Amazonas Chagas-Junior 1 & Renner L. C. Baptista

Laboratrio de Aracnologia, Departamento de Invertebrados, Museu Nacional. Quinta da Boa Vista, So Cristvo,
20940-040 Rio de Janeiro, Rio de Janeiro, Brasil.
2
Laboratrio de Diversidade de Aracndeos, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de
Janeiro. Avenida Carlos Chagas Filho 373, Cidade Universitria, 21941-902 Rio de Janeiro, Rio de Janeiro, Brasil.
3
Corresponding author. E-mail: [email protected]

ABSTRACT. A revision of the currently known distribution of Onychophora (velvet worms) in Brazil is presented. Twentyfour morphospecies (half of them undescribed) belonging to four genera of the Peripatidae (Peripatus Guilding, 1826,
Epiperipatus Clark, 1913, Macroperipatus Clark, 1913 and Oroperipatus Cockerell, 1908) are recorded. This high number
of unidentified morphospecies is due to three major reasons: the poor quality of the characters used in species-level
identification, the inadequate specimen conservation in ethanol, and, in most cases, the lack of additional specimens for
analysis of intra- and inter-specific variation. The morphological characters currently used to separate Peripatidae species present a large degree of variation and may not be sufficient to allow accurate identification. There are records of
these animals from 16 states, located in four political regions of the country (North, Center-West, Northeast and Southeast). Oroperipatus balzani is recorded for the first time from Brazil. The southernmost record for Onychophora is Itacuru
island, Mangaratiba, Rio de Janeiro State (E. edwardsii) and the northernmost is Vila Tepequm, Amajari, Roraima State
(Epiperipatus sp. 4). Considering the large territorial area of Brazil and the paucity of localities where Onychophora have
been collected, we may expect a total diversity of these animals much larger than the currently known one.
KEY WORDS. Brazil; distribution; Peripatidae; taxonomy; velvet worm.

Onychophora is an invertebrate phylum poorly known

by the layperson and even by most biologists. However, it is a
key taxon for the understanding of the evolution of
Panarthropoda (BRUSCA & BRUSCA 1990, MONGE-NJERA 1995).
Velvet worms are distributed in two large families, with around
200 species, and display a Gondwana distribution (MAYER 2007).
Species of Peripatidae are found in tropical areas (West Africa,
southern Mexico, Central and South America, Antilles and
Southeast Asia), whereas species of Peripatopsidae present a
southern temperate distribution (South Africa, Oceania and
Chile) (MONGE-NJERA 1995).
According to PECK (1975), 60 species and nine subspecies
of Peripatidae and one species of Peripatopsidae are known from
South and Central America. Species of Peripatidae are known
from southern Mexico and the Greater Antilles to southeastern Brazil. On the other hand, Peripatopsidae occurs only in
Chile. Up to date, four genera and 11 species have been recorded for Brazil (OLIVEIRA & WIELOCH 2005, SANTANA et al. 2008),
the largest species diversity in the Neotropics, notwithstanding the scanty work on the taxonomy of velvet worms. Most
Brazilian species display a limited distribution, as Macroperipatus
acacioi (Marcus & Marcus, 1955) (endemic from the Tripu Ecological Station in the city of Ouro Preto, state of Minas Gerais).
In contrast, Epiperipatus edwardsii (Blanchard, 1847) is found

from the Central America and the Lesser Antilles to southeastern Brazil (PECK 1975).
The first compilation on velvet worms distribution was
proposed in the seminal work of SEDGWICK (1888). Besides the
many new records of Onychophora, this was the first study to
propose the use of both internal and external characters in the
taxonomy of the group. Regarding the Neotropical species,
Sedgwick cited for the first time unidentified specimens of velvet worms from Brazil, collected in the vicinities of Santarm
and Breves, both cities in the state of Par.
The most important works about Onychophora are the
two monographs published in the beginning of the twentieth
century by BOUVIER (1905, 1907). Most of the knowledge on
taxonomy, biology, anatomy and distribution of Neotropical
velvet worms are still based on those works. Among the 32
species of Peripatidae considered in the first study (BOUVIER
1905), five are species from Brazil: Peripatus ohausi Bouvier,
1900, P. brasiliensis Bouvier, 1899, P. edwardsii, P. simoni Bouvier, 1898 and P. eiseni Wheeler, 1898.
Years later, CLARK (1913) presented a new classification
for the American species of Peripatus. This author proposed three
new subgenera, Plicatoperipatus Clark, 1913, Macroperipatus and
Epiperipatus, and a new genus, Metaperipatus Clark, 1913, to
accommodate the only know Peripatopsidae from the Ameri-

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554

cas. Soon after, CLARK (1915) discussed and presented a list of

hitherto known records of the species of Peripatidae and
Peripatopsidae from all over the world.
BRUES (1923) published the first map summarizing the geographical distribution of Onychophora throughout the world.
Besides a general world map, he elaborated maps depicting the
distribution of Onychophora in Australia, West Indies, Africa,
South and Central America. Although, this publication analyzed
only the distribution of the families and genera, but did not
present a detailed list of species and their records.
MARCUS (1937) described a new species, Peripatus (Epiperipatus) evelinae Marcus, 1937, from central Brazil. The new species presented a large number of lobopodes (32 and 34), an unseen characteristic in Brazilian velvet worms at that time. Four
years later, CARVALHO (1941) described Peripatus (Peripatus) heloisae
Carvalho, 1941, from Mato Grosso, also from central Brazil. This
species has a distinctive white spot on the head. CARVALHO (1942)
added new external and internal features for P. heloisae, based
on more than 600 additional specimens he collected. MARCUS &
MARCUS (1955) described a new species, Peripatus acacioi Marcus
& Marcus, 1955, from Tripu, in the vicinities of the city of Ouro
Preto, state of Minas Gerais, also in Brazil. The shape of the cuticular papillae from the new species is similar to P. ohausi, however both species differ in color and presence of accessory teeth.
The first revisionary paper on Brazilian Onychophora was
published by FROEHLICH (1968). He studied a small collection of
Onychophora from the former Departamento de Zoologia,
Faculdade de Filosofia, Universidade de So Paulo, Brasil. Comments on three species (M. acacioi, P. evelinae and E. edwardsii)
were given, and a new species, Peripatus tucupi Froehlich, 1968,
was described from the state of Par.
The last summary of the Onychophora distribution was
published by PECK (1975), but limited to the Neotropical region. In that paper, he described a new genus and gave a revised and updated version of the key to American genera of
Onychophora of BOUVIER (1905), including the taxa proposed
by Clark and himself. He cited nine species, belonging to four
genera, collected in 14 different localities from Brazil.
Most papers on Brazilian Onychophora in the last decades dealt with biology and physiology of M. acacioi (e.g.,
CAMPIGLIA & LAVALLARD 1973, LAVALLARD & CAMPIGLIA 1973). Only
after the beginning of the current century, studies on taxonomy
and distribution of velvet worms began to be published again
in Brazil. VASCONCELLOS et al. (2004) recorded specimens of three
Peripatus spp. (possibly new) for the first time from the states
of Pernambuco and Alagoas. OLIVEIRA & WIELOCH (2005) described a new species of Macroperipatus, M. machadoi Oliveira &
Wieloch, 2005, from the state of Minas Gerais. VASCONCELLOS et
al. (2006) recorded Onychophora species from the Atlantic
Rainforest in four states from northeast Brazil (Cear,
Pernambuco, Alagoas and Bahia). The specimens were identified only to the generic level. Later, SANTANA et al. (2008) added
the first record of Onychophora from the state of Paraba, north-

ZOOLOGIA 26 (3): 553561, September, 2009

C. Sampaio-Costa et al.

eastern Brazil, represented by one specimen collected in the

Atlantic Rainforest.
Notwithstanding the above-cited papers, there is no comprehensive paper on the currently known geographical distribution of Onychophora in Brazil. This study presents a species
list of Brazilian velvet worms, based on literature, museum specimens and firsthand records. Besides, some characters used in
the identification of velvet worms are reexamined and the currently known distribution of Brazilian Onychophora is analyzed.

MATERIAL AND METHODS

The list below is based on the literature and the analysis
of 798 specimens deposited at the following institutions (acronyms and curators given between brackets): Museu Nacional,
Universidade Federal do Rio de Janeiro, Rio de Janeiro (MNRJ, A.
B. Kury); Museu de Zoologia, Universidade de So Paulo, So
Paulo (MZSP, R. Pinto-da-Rocha); Instituto Butantan, So Paulo
(IBSP, A. D. Brescovit); and Departamento de Zoologia,
Universidade Federal de Minas Gerais, Belo Horizonte (DZUFMG,
A. Wieloch). The identification of the specimens follows BOUVIER
(1905) for the most part, except for the species described after
the publication of this revisionary paper. The generic classification used in the species list follows PECK (1975). Species records
marked by an asterisk (*) are those of specimens that were not
examined; all information for them are derived from, the literature. Data on localities (distribution section) were gathered from
the gazetteers and locality directory of the Falling Rain internet
site (http://www.fallingrain.com/world/). The locality record for
each specimen was included in a database and afterwards exported to ARC-GIS for map elaboration.

TAXONOMY
List of Brazilian Onychophora

Peripatus Guilding, 1826

Peripatus evelinae Marcus, 1937
Type-material: BRAZIL, Minas Gerais/Gois: between Nova
Roma and Cana Brava, Blaser leg. (male lectotype, female allotype and two paratypes, MZSP).
Specimens examined: only types.
Distribution: Cerrado areas (dry, open savannas) in the
border of Minas Gerais and Gois states.

Peripatus heloisae Carvalho, 1941 (Fig. 1)

Type-material: BRAZIL, Mato Grosso: Santa Terezinha (formerly Barra do Tapirap): A.L. Carvalho leg. (male holotype,
MNRJ 0002; female paratype, MNRJ 0003).
Specimens examined: BRAZIL, Tocantins: Miracema do
Tocantins: IX.2001, R. Bertani leg. (3 specimens, IBSP); Mato
Grosso: Santa Terezinha: A.L. Carvalho leg. (male holotype,
MNRJ 0002); (female paratype, MNRJ 0003); XI.1939, A.L.
Carvalho leg. (594 specimens, MNRJ 0001).

Brazilian species of Onychophora with notes on their taxonomy and distribution

555

Figures 1-4. Brazilian velvet worms: (1) Peripatus heloisae, adult; (2) Macroperipatus acacioi, adult; (3) Epiperipatus edwardsii, adult female
and juvenile; (4) Peripatus sp. 2.

Distribution: Cerrado areas from the states of Mato

Grosso and Tocantins.

Peripatus sp. 1
Specimens examined: BRAZIL, Cear: Pacoti: Macio de
Baturit: 22.X.1998, A. Percequilo leg. (1 specimen, MZSP).
Distribution: Cear State.

Peripatus sp. 2 (Figs 4 and 5)

Estao Biolgica Mata do Sossego, 7-19.III.1999, U. Caramaschi

leg., R.N. Feio, H. Niemeyer & M.F. Napoli leg. (1 spec., MNRJ
0012); Esprito Santo: Colatina: Rio So Jos (1 specimen, MZSP).
Distribution: Par, Tocantins, Minas Gerais and Esprito
Santo states.

Peripatus sp. 4
Specimens examined: B RAZIL , Bahia: Porto Seguro:
Trancoso, 19.VII.2005, A. Chagas Jr, B. Segal & E.G. Vasconcelos
leg. (1 specimen, MNRJ 0017).
Distribution: Bahia State.

Specimens examined: BRAZIL, Tocantins: Porto Nacional:

Luzimangue, 13.VII.2007, A. Chagas Jr, A. Giupponi, A. Kury
& A. Prez leg. (1 specimen, MNRJ 0030); Parque Estadual do
Lajeado: 11.VII.2007, A. Chagas Jr, A. Giupponi, A. Kury & A.
Prez leg. (1 specimen, MNRJ 0031); Lagoa das Confuses:
Caverna da Igreja, 08.VII.2007, A. Chagas Jr, A. Giupponi, A.
Kury & A. Prez leg. (2 specimens, MNRJ 0032, 0033).
Distribution: Tocantins State.

Specimens examined: BRAZIL, Bahia: Porto Seguro: Reserva

Vera Cruz Florestal, 2.XII.1994, O. Marques leg. (1 specimen,
MNRJ 0011).
Distribution: Bahia State.

Peripatus sp. 3

Peripatus sp. 6

Specimens examined: BRAZIL, Par: Conceio do Araguaia:

26.VII.1988, R. Pinto-da-Rocha leg. (1 specimen, MZSP); Tocantins:
Palmas: 23.II.2003, (1 specimen, IBSP); Minas Gerais: Simonsia:

Specimens examined: BRAZIL, Mato Grosso: Alto Araguaia:

04.VI.1994, J.M. Marins leg. (1 specimen, MZSP).
Distribution: Mato Grosso State.

Peripatus sp. 5

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C. Sampaio-Costa et al.

Epiperipatus Clark, 1913

Epiperipatus brasiliensis (Bouvier, 1899)
Type-material: BRAZIL, Par: Santarm: Wickham leg. (4
syntypes, MNHN, BMNH)*.
Specimens examined: BRAZIL , Maranho: Estreito: Rio
Farinha, Cachoeira da Usina, 7-14.I.2001, F.C. Straube leg. (1
specimen, MNRJ 0013); Cear: Crato, R. Alencar leg. (1 specimen, MNRJ 0022).
Distribution: Panama, Venezuela and Brazil (Par, Maranho and Cear states).

Epiperipatus edwardsii (Blanchard, 1847) (Fig. 3)

Type-material: FRENCH GUIANA, Cayenne: 3 mi from mouth
Approuage river, Lacordaire (female holotype, MNHN)*.
Specimens examined: BRAZIL, Par: Abaetetuba: Piratuba,
1938, A.L. Carvalho leg. (13 males, 11 females, MNRJ 0028);
Esprito Santo: Rio Doce, Porto Cachoeiro [On the specimen
label, it is written Porto Cachoeiro, Rio Doce. Porto
Cachoeiro is the former name of Santa Leopoldina, a city located around 44 km from Vitria. However, the Doce river is
almost 100 km away from Santa Leopoldina. So, the precise
locality can not be determined, due to the discrepancy in the
label], E. Garbe leg. (1 specimen, MZSP 483); Pinheiros: Reserva
Biolgica Crrego do Veado, 21-23.X.2005, Expedio ESFA
leg. (2 specimens, MNRJ 0023, 0027); idem, 22.X.2005, A.P.L.
Giupponi, et al. leg. (1 specimen, MNRJ 0026); Linhares:
Floresta Nacional de Goytacazes, 25.X.2005, A.P.L. Giupponi,
A. Kury, T. Bernab, T. Souza leg. (1 specimen, MNRJ 0020);
idem, 26.X.2005, A.P.L. Giupponi, A. Kury, T. Bernab leg. (2
specimens, MNRJ 0018); idem, Goytacazes, 13.X.1957, M. Rosa
leg. (3 males, 5 females, MNRJ 0029); So Mateus: Reserva
Florestal da Companhia do Vale do Rio Doce, VII.1998, A.J.
Santos & G. Machado leg. (3 specimens, MZSP 03709); Santa
Teresa: 2005, A.P.L. Giupponi leg. (1 mature and 1 young specimen, MNRJ 0034); idem, Estao Biolgica Santa Lcia, 2830.IV.2004, A.P.L. Giupponi, M.P Milleri & T. Souza leg. (2
specimens, MNRJ 0015); Rio de Janeiro: Mangaratiba: Ilha de
Itacuru, guas Lindas, I.1973, B. Prazeres leg. (1 male, MNRJ
0006).
Distribution: Panama, Trinidad & Tobago, Venezuela,
French Guiana, Surinam, Colombia and Brazil (Par, Esprito
Santo and Rio de Janeiro states).
Remarks: the only specimen known from Itacuru, Rio
de Janeiro, is dubiously included in E. edwardsii. It is in bad
state, and we have not been able to find any character to separate it from the other specimens of E. edwardsii.

Epiperipatus simoni (Bouvier, 1898)

Type-material: VENEZUELA, Distrito Federal, Caracas, E.
Simon leg. (female holotype, MNHN)*.
Specimens examined: BRAZIL, Rio de Janeiro: Barra de So
Joo, Morro de So Joo, 21-24.III.2003, Expedio Arachn (2
specimens, MNRJ 0014).

ZOOLOGIA 26 (3): 553561, September, 2009

Distribution: Venezuela and Brazil (Par* and Rio de

Janeiro states).
Remarks: the two specimens from Barra de So Joo are
very similar to the redescription and illustration of the type of
E. simoni presented by BOUVIER (1905). These two specimens
have been dubiously included in E. simoni, but they differ in
minor details as the presence or absence of frontal organs and
the number of small papillae between the primary papilla basis (PPB). The specimens from Barra de So Joo do not present
frontal organs and have 1-7 small papillae between the PPB,
whereas E. simoni presents frontal organs and 1-4 small papillae between the PPB, which may prove to be of specific value
following further studies.

Epiperipatus tucupi Froehlich, 1968

Type-material: BRAZIL, Par: 1939, E.C. Lima leg. (female
holotype and paratype, MZSP).
Specimens examined: only the types.
Distribution: Par State.

Epiperipatus sp. 1 (Figs 6-8)

Specimens examined: BRAZIL: Alagoas: Murici, Estao
Ecolgica de Murici, Nancy (1 specimen, MNRJ 0035); Murici,
Mata da Bananeira, 21.VII.1995, L.M. Freitas & M.S. Natali (1
specimen, DZUFMG-ONY 0008); Murici, Mata da Bananeira,
21.VII.1995, L.M. Freitas & M.S. Natali (1 specimen, DZUFMGONY0009).
Distribution: Alagoas State.

Epiperipatus sp. 2
Specimens examined: BRAZIL , Esprito Santo: Cariacica,
Reserva Florestal Duas Bocas, X.2004, A.L.P. Giupponi, M.P
Milleri & T. Souza leg. (1 male and 1 female, MNRJ 0016); Reserva
Florestal Duas Bocas, I.1987, R.L.C. Baptista leg. (1 specimen,
MNRJ 0010).
Distribution: Esprito Santo State.

Epiperipatus sp. 3
Specimens examined: B RAZIL , Amazonas: 1937, A.L.
Carvalho leg. (1 specimen, MNRJ 0024).
Distribution: Amazonas State.

Epiperipatus sp. 4
Specimens examined: B RAZIL , Roraima: Tepequm,
11.XII.1994, E.O.S. Sueliano leg. (1 specimen, MZSP).
Distribution: Roraima State.

Epiperipatus sp. 5
Specimens examined: BRAZIL, Amap: Territrio Federal
do Amap, 18.VI.1966, Froehlich leg. (1 specimen, MZSP);
Ferreira Gomes, Stio Otimari, km 322 da Br. 156, 20.X.2007,
J.F. Cludio & P. Magno leg. (1 male and 7 females, MNRJ 0036);
Porto Grande, Mata do Areal, 21.X.2006, J.F. Cludio & P. Magno
leg. (4 specimens, MNRJ 0037).
Distribution: Amap State.

Brazilian species of Onychophora with notes on their taxonomy and distribution

557

Figures 5-8. (5) Integument of Brazilian Peripatidae: Peripatus sp. 2, detail of the laterodorsal integument; (6) Epiperipatus sp. 1, dorsal
view of the disposition of PPB and distance among the folds; (7) Epiperipatus sp. 1, detail of the PPB; (8) Epiperipatus sp. 1, the PPB
extent. (PPB) Primary papilla basis.

Macroperipatus Clark, 1913

Macroperipatus acacioi (Marcus & Marcus, 1955) (Fig. 2)
Type-material: BRAZIL, Minas Gerais: Ouro Preto, Tripu,
17.X.1947, O. Schubart leg. (Holotype male, MZSP); idem
(Paratype female, MZSP); A. Costa (7 specimens, syntypes, MZSP).
Specimens examined: BRAZIL, Minas Gerais: Ouro Preto,
Tripu, 17.X.1947, O. Schubart leg. (Holotype male, MZSP); Ouro
Preto, Tripu, 17.X.1947, O. Schubart leg. (Paratype female,
MZSP); Ouro Preto, Tripu, A. Costa leg. (7 specimens, syntypes,
MZSP); Ouro Preto, Tripu, 12.VII.1961, G. Simes & A. Costa
leg. (1 specimen, MNRJ 0004); Ouro Preto, Tripu, VII.1961, A.
Costa & J.L.A. Feio leg. (6 specimens, MNRJ 0005); Ouro Preto,
J. Moojen leg. (4 specimens, MNRJ 0038); Ouro Preto, Tripu,
IV.2008, I. Oliveira leg. (4 specimens, MNRJ 0039).
Distribution: Minas Gerais State.

* Macroperipatus geayi (Bouvier, 1899)

Type-material: FRENCH G UIANA : Carsevenne Contest

Franco-Brsilien (now Amap State), 1898, F. Geay leg. (female

type, MNHN)*.
Distribution: Costa Rica, Panama, French Guiana, Colombia and Brazil (Amap State).

Macroperipatus machadoi Oliveira & Wieloch, 2005

Type-material: BRAZIL, Minas Gerais: Caratinga, Reserva Particular do Patrimnio Natural Feliciano Miguel Abdala, VII.1984
(Holotype female, DZUFMG-ONY 0001); idem, 21.XII.2001, J.C.
Silva-Jnior leg. (male, DZUFMG-ONY 0002); idem, 27.VII.2000,
J.C. Silva-Jnior leg. (male, DZUFMG-ONY 0003).
Specimens examined: BRAZIL, Minas Gerais: Caratinga, Reserva
Particular do Patrimnio Natural Feliciano Miguel Abdala, VII.1984
(Holotype female, DZUFMG-ONY 0001); Caratinga, Reserva Particular do Patrimnio Natural Feliciano Miguel Abdala, 21.XII.2001,
J.C. Silva-Jnior leg. (male, DZUFMG-ONY 0002); Caratinga,
Reserva Particular do Patrimnio Natural Feliciano Miguel Abdala,
27.VII.2000, J.C. Silva-Jnior leg. (male, DZUFMG-ONY 0003).
Distribution: Minas Gerais State.

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C. Sampaio-Costa et al.

* Macroperipatus ohausi (Bouvier, 1900)

Type-material: B R A Z I L , Rio de Janeiro: Petrpolis,
16.VIII.1898, Dr. Ohaus (two female types, ZMH and MNHN)*.
Distribution: Rio de Janeiro State.
Remarks: the additional specimen from MZSP examined
by READ (1988a) was not located in that collection (R. Pinto-daRocha, Universidade de So Paulo, pers. comm.). Following information from the collector of the MZSP specimen (C. A.
Brando, Museu de Zoologia de So Paulo pers. comm.), we tried
to collect new specimens on the precise collecting site. However, we have not been successful, as the site is located at an area
of urban sprawl in Petrpolis. So, M. ohausi has not been captured again since 1981, despite our efforts to collect it.

Macroperipatus sp. 1
Specimens examined: BRAZIL, Pernambuco: Recife, Dois
Irmos, O. Schubart leg. (1 specimen, female, MNRJ 0025)
Distribution: Pernambuco State.

Oroperipatus Cockerell, 1908

Oroperipatus balzani (Camerano, 1897)
Type-material: BOLIVIA, Coroico and Chulumani provinces
(two male types, MZT and MHNG)*
Specimens examined: BRAZIL, Acre: Marechal Thaumaturgo,
REAJ, 24.VIII.94, A.J. Cardoso leg. (1 specimen, MNRJ 0021).
Distribution: Bolivia and Brazil (Acre State).

*Oroperipatus eiseni (Wheeler, 1898)

Type-material: Mexico, Nayarit, Tepic (87 types, BMNH
and MNHN)*
Distribution: Mexico, Panama and Brazil (Amazonas State).

DISCUSSION
In this study, 24 morphospecies of Onychophora are recorded from Brazil, 12 of them corresponding to described species (see taxonomic list above), distributed in four genera: Peripatus, Epiperipatus, Macroperipatus, and Oroperipatus. Up to now, O.
balzani was known only from Bolivia. It is recorded for the first
time from Brazil in this paper. The Brazilian records of M. geayi,
M. ohausi and O. eiseni are based only on the literature. The other
12 morphospecies are not ascribed to any described species. This
high number of unidentified morphospecies is due to three
major reasons: the poor quality of the characters used in specieslevel identification, the inadequate specimen conservation in
ethanol, and, in most cases, the lack of additional specimens for
analysis of intra- and inter-specific variation.
Since BOUVIERs monograph on Onychophora (1905), few
additional characters have been used in the taxonomy of
Peripatidae. One character used by Bouvier to separate
Peripatidae into two large groups, Peripates andicoles and
Peripates carabes, is the number of papillae in the foot of
the lobopods IV-V. The Peripates andicoles (now included in
Oroperipatus) present four or more papillae in the foot of

ZOOLOGIA 26 (3): 553561, September, 2009

lobopods IV-V. On the other hand, Peripates carabes (now

included in Peripatus, Epiperipatus and Macroperipatus) present
only three papillae in those feet. This character was proposed
by BOUVIER (1905) and is well known in the literature.
The diagnosis of the three genera included in the Peripates
carabes is difficult and not clearly stated in the current taxonomic literature. After READ (1988a), the apparent lack of good
specific characters and the great uniformity among genera within
the Peripatidae, particularly the large Caribbean group, has been
the great obstacle to taxonomic progress. In the same paper,
she revised the morphological characters used to separate genera in Peripatidae through SEM examination, and pointed to
great variability in the integument characters. She remarked that
the degree of body contraction of one specimen may greatly
affect the appearance of its integument. A distended specimen
shows a very different pattern in the shape and disposition of
the dorsal papillae in comparison to a contracted one, for example. We also observed a large variation in integument characters, as color pattern and shape and distribution of dorsal papillae. Most of this variation may probably be explained by the use
of different methods to fixing and preserving the specimens, as
well as by the variable time-span they are kept in collections.
Following the papers by BOUVIER (1905), CLARK (1913) and
PECK (1975), the shape of the basis of the primary papillae, called
papilles principales by Bouvier, and the disposition of the dorsal papillae on the folds allow for the separation among Peripatus, Epiperipatus and Macroperipatus. However, there is variation
in the shape and disposition of the papillae even within one
genus, as for example in Peripatus. The primary papilla bases
(PPB from now on figure 5) of Peripatus are round and placed far
apart from each other. Epiperipatus also has round PPB (Figs 6-8),
but they are close together. Finally, Macroperipatus presents PPB
almost square and placed not far apart. As remarked above, the
fixing method and preservation period may have a substantial
effect upon the shape and disposition of PPB. In several of the
12 unidentified morphospecies in our list, it was easy to find
specimens presenting either papillae closely spaced or placed
farther away, as for example in Peripatus sp. 3 and Peripatus sp. 2.
The diagnostic character for Macroperipatus is the square
PPB (BOUVIER 1905, CLARK 1913, PECK 1975), as found in M.
torquatus (Kennel, 1883) (type-species), M. geayi, M. valerioi
Morera-Brenes & Lon, 1986 and M. perrieri (Bouvier, 1899).
On the other hand, M. acacioi and M. machadoi have PPB varying from subquadrate to round, but never square PPB. We have
not examined specimens from M. ohausi, the other species of
Macroperipatus from Brazil, but READ (1988a) noted that it also
does not have the square basis. According to our results and
also those from READ (1988a), the square PPB seems to be restricted to the Macroperipatus species from Central America and
northern South America. The species from southeast Brazil bear
PPB similar to the ones found in Peripatus or Epiperipatus.
An additional character used to separate velvet worm
species is the contrasting color pattern (for example, stripes,

Brazilian species of Onychophora with notes on their taxonomy and distribution

lozenges etc.) over and around the median dorsal line. The
pattern of light lozenges over a darker background is found in
at least three Epiperipatus species (E. trinidadensis (Sedgwick,
1888), E. broadwayi (Clark, 1913) and E. edwardsii) and also in
M. acacioi. In contrast, the lozenges are not clearly seen in some
E. edwardsii specimens from Esprito Santo State, and may even
not be noticed at all. Most Brazilian Onychophora have brownish to purple-brown hues, often presenting a darker, thin, stripe
over the median dorsal line. P. heloisae bears a contrasting whitish spot just behind the head.
Among the nine Epiperipatus morphospecies we found,
E. edwardsii and E. brasiliensis are easily identified. We were not
able to examine any E. imthurmi specimen. Regarding the other
Epiperipatus morphospecies, the analysis of the outer morphological characters were not enough to allow an accurate identification. For example, Epiperipatus sp. 5 has the leg number
and irregular papillae disposition found in E. imthurmi, but it
differs in coloration. Epiperipatus sp. 5 is dark purple, whereas
E. imthurmi may present colors varying from orange to brown.
The only unidentified morphospecies of Macroperipatus
seems to be close to M. ohausi, but it is found only in Pernambuco State, northeastern Brazil, while M. ohausi is known only
from Rio de Janeiro State, southeastern Brazil. Both species
present jaw blades with an inner accessory tooth and well-developed frontal organs. Macroperipatus sp. 1, though, displays
a row of 11 teeth placed after the accessory tooth of the inner
jaw, absence of secondary papillae placed laterally to the primary ones, and 30 pairs of legs, contrasting to the 10 teeth,
presence of such secondary papillae and 27-28 pairs of legs
found in M. ohausi (after BOUVIER 1905). Summing up, the gap
found in the morphological characters and the distance between localities indicate that M. ohausi and Macroperipatus sp.
1 are probably distinct species.
As shown above, the morphological characters currently
used to separate Peripatidae species present a large degree of
variation and may not be sufficient to allow accurate identification. The use of characters of inner structures, as number of
teeth in the jaws, and SEM may yield good results when applied to Brazilian Onychophora, as has been noticed by READ
(1988 a, b) and MAYER (2007) in other countries.
Most papers on the distribution of Onychophora are
worldwide-based, and do not present detailed records for each
country. Considering Brazilian Onychophora, most records are
restricted to the description of species. Also, many new records
have been presented in abstracts of zoology meetings, including new but invalid scientific names.
During our studies, we examined specimens of velvet
worms from twelve of the thirteen Brazilian states already cited
in the literature (Amazonas, Par, Cear, Pernambuco, Alagoas,
Bahia, Tocantins, Gois, Mato Grosso, Minas Gerais, Esprito
Santo and Rio de Janeiro). The exception is Paraba, as we have
not been able to examine upon any specimen from it. Besides,
we added first-time records for three states (Roraima, Acre and

559

Maranho) and 27 new localities from Brazil. Summing up,

Onychophora are currently known from 16 states, from all
Brazilian regions, except the South. The southernmost record
for Onychophora is Itacuru island, Mangaratiba, Rio de
Janeiro State (E. edwardsii) and the northernmost is Vila
Tepequm, Amajari, Roraima State (Epiperipatus sp. 4) (Fig. 9).
Most described Brazilian species are certainly endemic,
as they are known only from their type-locality (e.g., E. tucupi,
M. acacioi, M. ohausi, M. machadoi and P. evelinae). Some species are known from just one additional collection outside the
type-locality (e.g., P. heloisae, O. eiseni and O. balzani), but others have what seems to be wide distributions (e.g., E. edwardsii,
E. simoni and E. brasiliensis). Epiperipatus edwardsii is the most
widespread species in Brazil, but most records are from Par
and Esprito Santo states.
Species of Oroperipatus are known only from the state of
Acre, northwestern Brazil. Macroperipatus spp. are found in
several southeastern states and also from an isolated record
from Pernambuco, northeastern Brazil. In contrast, species of
Epiperipatus and Peripatus are found on a wide geographical area
covering most states in Brazil (Fig. 9).
The 24 here recognized morphospecies of Peripatidae represent a substantial increase over the 11 species cited by OLIVEIRA
& WIELOCH (2005) and SANTANA et al. (2008). Also, the known
distribution of velvet worms now covers most Brazilian states,
with the exception of the South region. Considering the large
territorial area of Brazil and the paucity of localities where
Onychophora have been collected, we may expect a total diversity of these animals much larger than the currently known one.
As we pointed out before, the method of preservation of
the specimens exerts a large effect over the characters currently
used in Peripatidae taxonomy. Most specimens kept in museums do not allow a precise identification but they are still useful as mementos of the distribution of velvet worms and pinpoints for future collection. Taking in consideration the apparently limited capacity of dispersal of most velvet worms, additional collection effort is badly needed, as many of the old
localities are under intense human pressure and extinction
looms at the corner.

ACKNOWLEDGMENTS
We thank Ricardo Pinto-da-Rocha (MZSP) and Ivo S. de
Oliveira (MNRJ/UFMG), for loaning specimens used in this study.
Eduardo Vasconcelos, for the precious informations on the
Onychophora from MZSP collection. To Elivaldo de Lima for
SEM operation at the Center for Scanning Electron Microscopy
of Museu Nacional, UFRJ. The establishment of this Center was
made possible by a grant from Cenpes/Petrobras, and is part of
the companys Thematic Network for Marine Environmental
Monitoring. Glauco Machado, Rogrio Bertani, Cleide Mendona
and Antonio Brescovit, for the donation of specimens to the
MNRJ collection. Carlos R.F. Brando (MZSP) for the precious
information on the locality in Petrpolis of M. ohausi and Adriano

ZOOLOGIA 26 (3): 553561, September, 2009

560

C. Sampaio-Costa et al.

Figure 9. Map showing the known distribution of the 24 morphospecies of Onychophora from Brazil.

Kury for helpful suggestions and support to this work. Rogrio

Bertani and Abel P. Gonzlez for the photos. CNPq and FAPERJ,
for the grants given to the two first authors.

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Submitted: 02.X.2008; Accepted: 25.VIII.2009.

Editorial responsibility: Gabriel Mejdalani

ZOOLOGIA 26 (3): 553561, September, 2009