S. B. Peck.
A Review of the New World Onychophora, with the Description of a New Cavernicolous Genus and Species from Jamaica.
Psyche 82:341-358, 1975.

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A REVIEW OF THE NEW WORLD ONYCHOPHORA
WITH THE DESCRIPTION OF A
NEW CAVERNICOWUS GENUS
AND SPECIES FROM JAMAICA*
BY STEWART B. PECK
Department of Biology, Carleton University Ottawa, Ontario KIS 5B6, Canada
Few animals are as avidly sought and as rarely found by inverte- brate zoologists in the American tropics as are ony~ho~horans. Yet, attempting to determine any specimens which one is fortunate enough to find can be a frustrating experience because of the scat- tered and incomplete literature.
In December 1972, while engaged in a survey of the invertebrate cave fauna of Jamaica, I discovered an eyeless and depigmented onychophoran. At that time, I knew that the only other known species olf cave-adapted (troglobitic) onychophoran, Peripatopsis alba Lawrence 1931, was from one cave in South Africa. To evaluate the status of the Jamaican troglolbitic species I undertook a review of the literature of New World Onychophora, and an examination of some 300 specimens in the collections of the Museum of Compara- tive Zoology (Harvard University) and the Institute of Jamaica. As a result, I have concluded that a summary of the literature on the systematics and distribution of New World onychophorans would be of interest, with the description of the new genus and species. General treatments of Onychophora morphology, anatomy, and biology are those of Cuenot (194g), Zacher (1933), Kaestner (1968), Barnes (1968) and the references cited in these and other invertebrate texts.
The only systematic treatment of the Onychophora of the world is that of Bouvier (1905, 1907). Sedgwick (1908) reviewed the classification but did not formally add ,to it. Cuenot (1949) and Brues, Melander and Carpenter (1954) characterize the families and subfamilies of the world. No work unites what is known of all New World species. Brues ( 191 I ) presented a key 'for some species from the Caribbean and surrounding mainlands, adapted from Bou- vier. Clark (1913a) revised the American species on a generic level, and ( 191 3b) gave an annotated list of species. Keys for the West Indian species are available only for the Islands of Hispaniola (Brues, 1935) and Jamaica (Arnett, 1961). dark and Zetek *Manuscript received by the editor July 10, 1975.



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342 Psyche
[September-December
(1946) gave a key for species known from the Canal Zone and adjoining regions.
Distribution has been discussed by many, including Brinck (1956), Brues (1923) and Clark (igi^a, 1g13b, igi^a, b). The circum- tropical distribution of the 'family Peripatidae and the generally south temperate distribution of the family Peripatopsidae have led several authors to speculate that these distributions have been achieved through a Gondwana or continental drift mechanism (for instance, Cuenot, I 949 ; Hutchinson, 1928). However, although continental drift may be a fact, onychophoran distributions have not necessarily been controlled by it alone, and active, dynamic biological factors were probably more important in forming the distributions (Brinck, 1956 ; Darlington, 1965).
Since the summary papers by Bouvier and Clark, several species have been described and others have been placed in synonymy. The following key to the genera and list of species unites this dispersed literature and hopefully will provide a basis for encouraging further studies.
Key to the families and genera of New World onychophorans Ia.
Sexual opening between last pair of legs (Fig. I) ; Peripatop- sidae, Peripatoidinae; south-temperate distribution in southern ................................................ and central Chile. Metaperipatus
ib. Sexual opening between penultimate pair of legs (Fig. 2) ; . . . .
Per~patidae, Per~patmae. ............................................................ 2 2a.
Nephridial tubercles (urinary papillae) of fourth and fifth legs indenting proximal margin of third creeping pad band on leg, usually dividing third pad into two segments (Fig. 3) ; four or five creeping pads; four foot papillae, 2 on each side of foot. .......................................................................... Oroperipatus Not the above combination; nephridial tubercles o,f fourth and fifth legs perhaps connected to third pad and indenting its prox- imal margin, but opening in space between third and fourth creeping pads, usually indenting fourth pad or rarely dividing fourth pad into two segments (Figs. 4, 7) ; usually four (rarely five or three) creeping pads (Fig. 4) ; three or four foot papil- lae. ................................................................................................ 3 Four foot papillae (three anterior and one posterior, or rarely two anterior and two posterior) (Fig. 4) ; El Salvador. ............ ............................................................................. Heteroperipatus Three foot papillae (two anterior and one posterior, Fig. 7). .. 4



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19751 PeckÌÔNe World Onychophora 3 43
Twenty-four transverse dorsal folds to each segment, only slightly distinct because of numerous anastomoses and irregu- larities in grooves which separate them; 37 to 41 pairs of legs; .............................................................. Jamaica. Plicatoperipatus
Twelve transverse dorsal 'folds to each segment, separated by continuous grooves except at level of legs where some grooves ...........................
usually bifurcate; 36 or fewer pairs of legs. 5
..........
Eyeless; 22 or 23 pairs of legs; Jamaica. Speleoperipatus
............................................ With eyes; 27 to 36 pairs of legs.
6
Primary papillae of dorsal surface with a quadrangular base separated by straight grooves parallel with axis of body, dorsal body folds apparently divided by grooves; accessory papillae usually small and few in number. .................... Macroperipatus Primary papillae of dorsal surface with a rounded base, dorsal body folds usually appearing undivided ; accessory papillae with diverse degrees of development. ................................................ 7 Primary papillae of dorsal surface with great size differences in all a,ges, some very predominant, cylindrical, the other conical, smaller, usually three of the smaller or accessory papillae be- tween two of the larger ones ; the primary. papillary tubercles separated by rather broad intervals where the accessory papillae occur; crural tubercles in male usually present on more than ........................................ two pregenital pairs of legs.
Peripatus
Primary papillae of dorsal surface all of one type; in medium or large specimens these papillae intergrading through all sizes, while in small specimens some are conspicuously larger, papillae closely set though with occasional, accessory papillae between them; crural tubercles in male present on only the two pre- ................................................ genital pairs of legs. Epiperipatus
find it difficult to separate the last two genera with certainty. -
They have been used commonly as subgenera of Peripatus sensu lato and this might be a better arrangement in view of their lack of external distinctness. A study of the integumentary papillae with a scanning electron microscope (Peck, in preparation) might help to clarify the utility of these characters in Peripatus and Epiperi- patus.
Froehlich (1962) noted the need for a thorough review and revision of the complicated taxonomy of the New World onycho- phorans.
This should be based on internal
as well as superficial
external characters. There is probably as much future work in the detection of synonyms as there is in finding and describing new
species. As long ago as 1913, Clark (1913b) stated that "it is



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344 Psyche [September-December
astonishingly easy to make mistakes in their determination." This
is still true, partly because no key to the species exists. It was beyond the intent of this study to prepare one. In the following list, references to Bouvier ( 1905, 1907) and 'Clark
( 19 I 3a, I 9 I 3b, I g I 5a) are not individually given because they apply to almost all species. Also, dates and literature citations to descriptions of Bouvier ( 1905, 1907) and earlier are not given, because they may be fo'und in these works. The word '(seen" indi- cates specimens from the Museum of Comparative Zoology and the Institute of Jamaica on which some data are based. All new records reported here are based on specimens of the Museum of Comparative Zoology. A history of synonymy of each species is not given because the intention of the list is only to report presently accepted species and their documented localities.
Family Peripatopsidae
Metaperipatus dark, I 9 I 3a
Type species. Peripatus blainvillei Blanchard, 1847. By original designation and by monotypy.
Distributed only in southern and central Chile. M. blainvillei ( Blanchard) .
Southern Chile. Loncoche ; Puc6n ;
Palguin ; San Carlos ; Chi106 Island ; Corral ; Volcin Villarica ; Enero, in Cordillera Peluda in Valdivia, Prov.; Contulmo, in Cordil- lera Nahuelbuta separating Mallerco and Arauco Provinces ; Buchoco valley between Lake Lanalhue and sea; south of Cafiete, 10 km from Contulmo. Bouvier, 1928 ; Claude- Joseph, I 927, 1928 ; Gravier and Fage, I 925 ; Johow, I g I I.
M. costesi (Gravier and Fage), 1925. Chile: Prov. Colchagua; Marga Marga near Santiago; "central Chile." Gravier and Fage, 1925.
M. umbrinus (Johow), 191 I. Chile, near Zapallar, on coast of Aconcagua Prov., at 32'33'20'' S. lat. Johow, I 9 I I. Family Peripatidae
Epiperipatus Clark, 19 I 3a
Type species. Peripatus edwardsii Blanchard, 1847. By original designation.
Distribution. From Santarem, Brazil, through French Guiana, Surinam, Guyana, Trinidad, Tobago, and Grenada and westward in South America to Central America and north through Nicaragua



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19751 Peck -New World Onychophora 345
to Guatemala and possibly British Honduras (Belize). E. barbouri ( Brues) , I g I I. Grenada : Grand Etang, 2000 f t. seen. Carriacou ( ? ) , seen. Tobago : Scarborough. B rues, I g 1 I, I 9 I 4 ; Clark, 1937.
E. biolleyi (Bouvier). Honduras: San Juan Pueblo. Brues, 1925. British Honduras: near Benque Viejo ( ?), seen. Costa Rica: "La Estrella" ; Parismina ; San Jose ; Surubres, near San Maeto ; fruit farm near Siquirres, 6.vi.1930, Waldeck, I, new record; Turrialba, 6.vii. 1956, C. Gans, 4, new record ; Turrialba, 25.vii-I 5.viii.1965, A Chickering, 6, new record; Talamanca, no other data, 4, new record; Orotina, 30 Aug. 1920, E. R. Dunn, 2 in hollow tree 6 ft. above ground, new record. Panama: El Ceremeiio. Brues, 191 I ;
Clark, 1937 ; Clark and Zetek, I 946 ; Picado, I 91 I .. E. brasiliensis brasiliensis ( Bouvier ) . Brazil : Santarim. Ven- ezuela : Merida. Bouvier, I 908.
E. brasiliensis vagans Brues, 1925. Canal Zone: Barro Colorado Island, seen; Las Cascadas, seen; Fort Sherman, seen; Rio Chinilla, seen; Balboa. Panama: San Pablo, seen; Rio Tapia, seen. Brues, I 925 ; Clark and Zetek, I 946.
E. edwardsi (Blanchard) . Brazil : Rio Guam& BeGm do Para ; Port0 Cachoeira, Rio Doce, Espirito Santo State. French Guiana: Cayenne. Surinam : Paramaribo. Trinidad ( ?) . Venezuela: Haute Sarare; Bas Sarare; Mirida; Val'encia; Caracas, seen. Colombia: Santa Marta Mts., seen; Cincinnati Coffee Plantation in Santa Marta Mts., seen. Panama: Panama Station: Dariin; El Cermefio; Balboa. Canal Zone: Barro Colorado Island, 3.viii.1925, C. W. Dodge, I, new record ; Barro Colorado Island, I 7- I g.iii. I 967, Pat- terson exp., 3, new record. Arn,ett, 1961 ; Bouvier, 1906; Brues, r 9 14, I 925 ; Clark and Zetek, I 946 ; Froehlich, I 968. E. evansi (Bouvier). Guyana: east bank of Demerara River. E. imthurmi ( Sclater) . Surinam : Paramaribo, seen. French Gui-
ana : Cayenne ; Haut Carsevenne. Guyana : Essequibo ; Georgetown (Demera,ra) ; Maccasseema on Pomeroon River; Hoorubea on De- merara River ; Dunoon ; Wakapa ; Issororo. Trinidad. Colombia :
Santa Marta, Cincinnati Coffee Plantation, 3000 ft., seen. Brues, 19 I 4, I 925 ; Clark, I 9 I 5 b ; Froehlich, I 968. E. isthmicola ( Bouvier) . Costa Rica : San Jose; Cachi ; Santa Clara, Colombiana Farm, seen ; Hamburg Farm, I .iii.1930, F. Nev- ermann, one on dry wood under loose bark, new record; Guiipiles, 7 Aug. 1920, E. R. Dunn, 4, new record; Navarro, 22 Aug. 1920, E. R. Dunn, 8, new record; Zent, 22-30 July 1920, E R. Dunn, 6, new record. Brues, 1925.




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346 Psyche
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E. lewisi Arnett, 1961. Jamaica: John Crow Mts., 10 mi. SW Priestmans River; 5 mi. SW Priestmans River, 1500 ft.; I mi. WSW Ecclesdown; all in Portland Parish, seen. E. nicaraguensis (Bouvier) . Nicaragua : Matagalpa ; San Benito. E. simoni
( Bouvier ) . Brazil : Para ; Breves, on Maraj6 Island. Venezuela : Caracas. Clark, I g I 3c ; Clark and Zetek, I 946. E. trinidadensis trinidadensis (Stuhlmann). Trinidad: Port of Spain, seen ; Maraval Valley, seen ; Laventille, seen ; Simla, I 5.iv. 1964, A. M. Chickering, I, new record; Simla, 1960, C. Carter, I, new record. Brues, 1914.
E. trinidadensis broadwayi Clark, I 9 I 3. Tobago, seen. Clark 1913C.
E. tucupi Froehlich, 1968. Brazil : Para. E. vespucci Brues, 1914. 'Colombia: Santa Marta Mts., Cincinnati Coffee Plantation, 2300 it., seen.
Heteroperipatus Zilch, I 954a
Type species. Heteroperipatus engelhardi Zilch, 1954a. By orig- inal designation and by monotypy.
Known only in El Salvador and Panama.
H. engelhardi Zilch, 1954. El Salvador: Finca El Carmen, Vol- can San Vicente, I 100-1300 m, in litter-filled grooves in coffee plantation.
H. clarki (Dunn), 1943. New combination, suggested by Zilch, I 9543- Panama : Azuero Peninsula,.
Macroperipatus Clark, I 9 I 3 a
Type species. Peripatus torquatus von Kennel, 1883. By original designation.
Distribution. Rio de Janeiro north to French Guiana, Guyana and Trinidad, west to Panama, and north to Veracruz, Mexico; and on Hispaniola and Jamaica.
M. acacioi Marcus and Marcus, 1955. Brazil: Minas Gerais, Our0 PrEto, I 100 m. Froehlich, I 968.
M. geayi (Bouvier). French Guiana; on the Brazilian border. Colombia: Sierra Nevada de Santa Marta, seen; Sierra Nevada de Santa Marta, Cincinnati Coffee Plantation, seen. Panama: La Chorrera. Canal Zone : Barro Colorado Island, seen ; Pedro Miguel ; Balboa. Costa Rica; Osa Peninsula, Rincon, 1o.viii. 1966, S. B. Peck, I under log at 400 ft., new record. Brues, 1925; Clark, 1913b ; Clark and Zetek, 1946.
M. guianensis (Evans). Guyana: east bank of 'Demerara River.



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19751 Peck - New World Onychophora 347
M. insularis insularis Clark, 1937. Haiti: between Jacmel and Tronin.
M. insularis clarki Arnett, 1961. Jamaica: Portland Parish, 5 mi. "SW Priestmans River, 1500 ft., seen.
M. ohausi (Bouvier). Brazil: near Rio de Janeiro; Petr6polis. M. perrieri (Bouvier) . Mexico : Veracruz. M. torquatus (von Kennel). Trinidad: Port of Spain, seen; Maraval Valley, seen. Brues, I 9 14.
Oroperipatus Cockerell, I 908
Type species.
Peripatus lankesteri Bouvier, 1899. Believed to be by subsequent designation of Clark, 1g13a. Original paper un- available.
Distribution. The Pacific slope of tropical America from Tepic, Mexico, southward through Colombia, Ecuador, and Peru to Sorata, Bolivia, and in western Brazil.
0. balzani (Camerano). Bolivia : Coroico and Chulumani states. 0. belli (Bouvier) . Ecuador: Durin, on Guayas River. 0. bimbergi ( Fuhrmann) , I 91 3.
Colombia : Amagatal, 900- 1800
m ; Guaduas, 800 m. Fuhrmann, 1914.
0. bluntschli Fuhrmann, 1915. Peru: Rio Samiria, 120 m. Zilch, 1954b.
0. cameranoi ( Bouvier ) . Ecuador : Cuenca; Sigsig. 0. corradoi (Camerano). Ecuador : Quito ; Balzar ; Gua~aquil. Panama : El Ceremefio. Canal Zone : Barro Colorado Island, seen ; Ancon. Venezuela: Falcon State, Pauji Acosta District, H. G. Kugler, 1.xi.1945, 5, new record. Brues, 1925; Clark, 19148, 1914b ; Clark and Zetek, 1946.
0. ecuadoriensis (Bouvier) . Ecuador : Bulim. 0. eiseni (Wheeler). Brazil : Rio Purus. Panama: El Cermefio ; Panama City. Mexico : Tepic. Clark and Zetek, 1946 ; Furrmann, 3914.
0. goudoti (Bouvier). Mexico.
0. intermedius ( Bouvier ) . Bolivia: Sorata. 0. koepckei Zilch, 1g54b. Peru: western side of Andes, 35 km from Olmos, toward Jakn, 1400 m.
0. lankesteri ( Bouvier) . Ecuador : Paramba, near Quito. 0. multipodes (Fuhrmann) , 191 3. Colombia, Rio Arnagh. Fuhr- mann, 1914.
0. omeyrus Marcus, 1952. Peru: San Jose de Lourdes, on Rio Chirinos, 1000 m; Sahuayaco, between Abancay and Marcos, Uru- bamba valley, 800 m. Zilch, 1954b.




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348 Psyche - [September-December
0. ( ?) peruanus (Grube) .
Peru. Marcus, I 952 ; Zilch, I 954b.
0. peruvianus Brues, I 9 I 7. Peru : Tabaconis, near Huancabamba,. 6000 ft., seen. Zilch, 1g54b.
0. quitensis (Schmarda) . Ecuador : high regions. 0. soratanus ( Bouvier ) . Bolivia : Sorata. 0. tu berculatus ( Bouvier ) . Colombia : Popayin. 0. weyrauchi Marcus, 1952. Peru: Yurac on Rio Aguaytia, 300" m. Zilch, 1954b.
Peripatus Guilding, I 825, sensu stricto of Clark, I 9 I 3a Type species. Peripatus jdif ormis Guilding, I 825. By monotypy- Distribution. West Indian Islands of Jamaica, Haiti, Puerto Rico, Vieques, St. Thomas, Antigua, Monserrat, Barbados, Guade- loupe, Dominica, and St. Vincent, and in South America in Brasil and from the mountains of western Venezuela westward to Colom- bia and north of Panama and Costa Rica.
P. antiguensis Bouvier. Antigua: Barlar, near Warburton. Mon- serrat. Clark, I 929.
P. bavayi Bouvier. Guadeloupe.
P. bouvieri Fuhrmann, 1913. Colombia: Boca del Monte, near Bogota; Cundinamarca Dept., above Aguadita, wet forest with ferns, 26.iii.1967, W. Brown, I, new record. Fuhrmann, 1914. P. brolemanni Bouvier. Venezuela: Tovar, near Merida; Raxto Casselo ; Puerto Cabello.
P. dominicae dominicae Pollard. Dominica : Laudat ; Prince Ru- pert ; Roseau, seen.
P. dominicae barbadensis Froehlich, 1962. Barbados: St. John, Codrington College.
P. dominicae basilensis Brues, I 935. Haiti : Mount Basil, 4000 ft., seen. Dominican Republic: Mt. Diego de Ocampo, 3500-4000 ft., seen; north slope Lorna Rucilla, 8000 ft., seen; north of Lorna Rucilla, 6000-7000 ft., seen; Constanza, 3000-4000 'ft., seen ; Vic Valle Nuevo, 7000 ft., seen. Brues, 1935 ; 1939. P. dominicae darlingtoni Brues, 1935. Haiti: Massif de la Hotte, between Camp-Perrin and Mafin, 3000 ft., seen. P. dominicae haitensis Brues, 1913. Haiti: Furcy (type locality) ; La Visite, La Selle Plateau, seen. Brues, 1935. P. dominicae lachauxensis Brues, I 935. Haiti : fitang Lachaux, Massif de la Hotte, under 1000 ft., seen. P. evelinae (Marcus), 1937. Brazil : between Canra Brava and Nova Roma, near border of Goiis and Minas Gerais. Froehlich, I 968.




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19751 Peck -New World Unychophora 349
P. heloisae Carvalho, 1941. Brasil : Rio Tapirape, near confluence with Rio Araguaia. Carvalho, 1942.
P. julif ormis Guilding. St. Vincent.
P. juliformis danicus Bouvier. Virgin Islands: St. Thomas. St. Croix. St. John: along trail from Centerline Road to Cinnamon Bay, in litter in dry north-facing gully, 3.1.1967, S. Peck, I, new record. Brues, 1925 ; Clark, 1937.
P. juanensis Bouvier.
Puerto Rico : Arecibo ; Utuado ; Vieques Island ; El Yunque, May 1938, Darlington, 3000 ft., I, new record ; Manati, 2.vi.1943, F. Mora, Indoors, I, new record ; Toro Negro Forest Preserve, trail up Cerro Dona Juana, 28.xii.1~66, S. Peck, 6 on Fomes woody fungi bract on log on ground, 600 m, new record, specimens lost. Clark, I 9 I 3b ; I 9 I 30. P. manni Brues, 1913. Haiti: Furcy; La Visite, La Selle Plateau, 5000-7000 ft., seen. Brues, 1935.
P. ruber Fuhrmann, 1913. Panama : Chiriqui Prov., Lino, near Boquete, 4100-4500 ft. Costa Rica : Rancho Redondo. Guatemala : Puerto Barrios ( ?). The record of P. (Epiperipatus) biolleyi var. betheli Cockerel1 1913 from Puerto Barrios appears to be P. ruber according to Clark and Zetek ( 1946). Clark, 1941b; Fuhrmann, 1914-
P. sedgwicki Bouvier. Venezuela : Caracas, seen ; San Esteban ; La Moka ; Las Trincheras ; La Guayra.
P. swainsonae Cockerell. Jamaica : Bath ; Savanna la Mar ; local- ities in Hanover, Portland and Trelawny Parishes, from 400 to
I 500 feet, seen. Andrews, I 933 ; Arnett, 1961 ; Brues, 191 I ; Lynn, 1934.
Plicatoperipatus Clark, I 91 3a
Type species. Peripatus jamaicensis Grabham and Cockerell, 1892.
By original designation and by monotypy. Known only from Jamaica.
P. jamaicensis (Grabham and Cockerell). Jamaica: Portland, Manchester, St. Thomas, Trelawny, St. Ann, and Westmorland Parishes, seen. Andrews, I 933 ; Arnett, I 961 ; Brues, 19 I I ; Lynn, 1936.
Speleoperipatus new genus
Description. Peripatidae, Peripatinae, with 22 or 23 pairs of legs; three foot papillae (two anterior and one posterior) ; legs 'four and five with urinary papillae broadly attached to and indenting creeping pad three, pad four cre~cent~~haped; no fifth pad. Type species. Speleoperipatus speleus, new species.



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3 50 Psyche [September-December
Speleoperipatus speleus new species
Holotype deposited in invertebrate collections of the Museum of Comparative Zoology, Harvard University. Type locality: Jamaica, Clarendon Parish, Pedro River, Pedro Great Cave. Type data: 20 December 1972, S. B. Peck. Paratypes: all from same locality; one, 25 March 1973, R. Norton and R. Zimmerman (deposited in Peabody Museum, Yale University) ; two, I 7 August 1974, S. Peck and family, in Museum of Comparative Zoology. Description. Eyes absent, no trace remaining. Depigmented ; specimens in life pale, almost white, with a slight greenish-yellow cast (Fig. 8). Legs and "antennae" not appreciably elongated. Holotype 34 mm long, with 22 pairs of legs. Paratypes: One 27 mm Figures 1-6. 1. Location of genital opening in 6
Peripatopsidae, Peripa-
toidkae. 2. Location of genital opening in Peripatidae, Peripatinae. 3. Ventral surface of left fourth leg of Oroperipatus koepckel (after Zilch, 1954b) showing urinary papilla dividing the third band of the creeping pads, and four foot papillae
(fp). 4. Ventral side of right fourth leg of Heteroperifatus engelhardi (after Zilch, 1954a) showing urinary papilla between creeping pads three and four, indenting pad four, and three anterior and one posterior foot papillae (fp). 5. Outer blade of right mandible of Speleoperipatus spelaeus. 6. Inner blade of right mandible of S. spelaeus.




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