Rev. Biol Trop., 34(2): 277-282, 1986

Description of Macroperipatus valerioi n. Sp. fron Costa Rica, and cornrnents on the genus Macroperipatus (Onychophora: Peripatidae)

Bernal Morera-Brenes and María D. León Musco de Zoología. Universidad de Costa Rica

(Received: January 21, 1986)

Abstract: A Ma croperípatus va/eríoí new species, Morera-Brenes & León, is described from Río Damitas, 16 Km north of Puerto Quepos (90 34' N, 840 lO' W), Costa Rica. The inner jaw bears 2 accessory teeth, the first more developed and with 13 denticles. Outer jaw with only one accessory tooth. Fourth creeping pad of fourth and fifth lobopods is slender and curves itself around the nephridial tuberc1e, which remains free and direc1ed towards the right side of the third and fourth pads. Fourth pad complete or broken in a single indi­ Macroperipatusgeagi vidual. Uniform dorsal color brown in life. This species appears to be c10sely related to (Bouvier) from South America.

Clark (1913 a) raised the torquatus section altitude 600 m. Data: 22-January-1965: A. of Peripatus (sensu lato) to generic status as Wille & M. E. Bozzoli. Macroperipatus. Brues (191 1) published a key to that taxon, to wruch several species have Etimology: M. valerioi Morera & León, been added. Annotated species checklists can 1986 is dedicated to the Costa Rican natura­ be fo und in Clark (1913 b) and Peck (1975). list and arcachnologist Carlos E. Valerio. Other Iiterature pertinent to the genus: Bouvier (1899, 1905); Brues (1911, 1925); Clark Diagnosis: inner jaw with 2 accesory teeth, (l913a, 1913b, 1937); Clark and Zetek (1946); first more developed and with 13 dentic1es. Von Kennel (1885, 1886) and Marcus and Outer jaw: only one-accesory tooth (Fig. la, c). Marcus (1955). In fourth and fifth pairs of lopods, the fourth The biology of species of Macroperipatus is creeping pad is trun and twists around the uri­ practically unknown, except for M. acacioi of nary tuberc1e, wruch is free and outerly bound. Minas Gerais (Brazil) (Campiglia et al. 1973, Fourth creeping pad complete or broken 1975, 1982, Lavallard et al. 1973, 1974, (Fig. 2a). 1975 a, b, c, Amaral �t al. 1980. Campiglia 1976, Description: Lavallard 1981) and a study of feeding habits 8.5 mm long, 0.5 mm max. height, 0.85 mm max. width, female, 34 pairs of M. torquatus which began recently (Read 1985). The systematics of Costa Rican ony­ of lobopods. Color (alive): uniform brown in dorse (A. Wille, 1985 , pers. com.). Preserved chophorans is currently under study by the sen­ in FAA: light beige, lighter ventrally. Eyes oval ior author and collaborators, and valerioi is M. (0.2 x 0.29 mm). Antennae 0.6 mm long, more the fi rst of several new species to be described. than 50 segments. No aparent frontal organs. Macroperipatus valerioi n. sp. Preserved �ith mouth evaginated ; lips: a pair of tuberc1es �lus superior and inferior ones (not Type: Museo de Insectos, Universidad de paired). qbrum with 9 dentic1es, the eighth bi­ Costa Rica. cuspidal (�ig. 1 b). Outer blade of jaws with one accesory ,ooth (Fig. la). Lobopods with 4 Locality: Costa Rica: Río Damitas, 16 km creeping pads, except last 3 pairs wruch are ru­ north of Puerto Quepos (90 34' N; 840 10' W), dimentary. AII feet with 2 anterior and one pos-

277 278 REVISTA DE BIOLOGIA TROPICAL

terior papillae, which have at least two bristles. accesory papillae of M geagi, as we fo und in Opening of coxal g1and long and slender. M valerioi. Clark (1939) says of M insularis Lobopods l.5-3 mm long and 0.18-0.36 mm that the less developed accessory papillae wide. Body segments with 12 dorsal skin folds outnumber primary papillae. This also occurs in each, 7 of which reach ventral side (the rest in­ M acacioi (Marcus & Marcus, 1955) and M complete). Primary and accessory papillae valerioi. This supports inclusion of the new spe­ projecting from square or oblong bases, with cies in Macroperipatus, as does the c1early quad­ traverse folds divided by shallow groves. Bases rangular base (of papillae) if compared with 0.07-0.43 mm long in each fold. Mid-dorsal Central American Ep ip eripatus biolleyi, E. line wide , made by a parallel sequen ce of bases isthmicola, and Peripatus ruber which have without papillae or with very small papillae, rounded bases, and with M geagi of Colombia. and not by a traverse crevice across folds. Line We could not examine types of M geagi and M chanalized between two papillae, primary and perrieri (currently studied by H. Ruhberg, Ham­ accesory (Fig. 3a) . Primary dorsal papillae burg Museum), but used Bouvier's (1899) uniserial and similar in size, like a round or drawings and a specimen of M geagi from Co­ conic breast, topped by a thin, almost cylindri­ lombia kindly provided by H. Ruhberg. In di s­ cal cone, with terminal bristles (Fig. 4a). Acces­ tribution, the quadrangular base of papillae of sory papillae conic, topped by a cone identical M valerioi is similar to that of M acacioi to that of a primary papillae (Fig. 4b). Size of (Marcus & Marcus 1955: Fig. 3d). Geograph­ cones is variable but smaller than primary ically, the c10set Macroperipatus are M. geagi cones. Rarely biserial (Fig. 2a, 4a). Primary (French Guiana, perhaps ranging to Panamá), papillae normally separated by one or several M Torcuatus (Trinidad) and M perrieri (Méxi­ secondary papillae, with no obvious pattern. co). Primary dorsal papillae of M. valerioi are Ventral fossae as usual. different from those of M perrieri because they lack terminal cones and are short and piramidal. Natural History: found below fallen log, M perrieri lacks accessory papillae. Accessory forest near Damitas river, in Premontane Rain papillae in M. valerioi are breast like (rounded Forest in Holdridge's (1978) system. or conical), and topped by a conic cylinder. This is similar in M geagi, where they are mark­ DISCUSSION kedly conical and topped by a long cone (Bouvier 1899). Mid-dorsal line is wide in M. Bouvier divided American onychophores as valerioi and is not a depression, both conditions "andicoles" or "cariiibes". The latter with 3 in contrast with M perrieri but similar to M papillae in lobopods, two anterior, one poste­ geagi, in which the line is chanelized between rior. Urinary papilla of lobopods N and V be­ two small papillae only , however (Bouvier, low third arc of creeping pads, deeply insert 1899). Pattern of papillae of oral tubercles the fourth. It was subdivided in three groups makes the mouth different from that of M corresponding to the currently accepted genera geagi. Jaws of M valerioi differ from those of Macroperipatus Clark, Epip eripatus Clark and M perrieri and M geagi (one accessory tooth. Pe ripatus (Guilding) (s. ste.). An eyeless genus Fig. Id), and from M torcuatus (second ac­ described recently, Sp eleoperipatus Peck, is cessory tooth more developed. Fig. 19)*. c10sely related to the latter two. These taxa are Pattern of creeping pad arcs different from tho­ a natural group also defined by the presence of se of M torcuatus and M. perrieri in which 12 folds per segmento P/icatoperipatus Clark there are signs of a fifth are (Fig. 2e) (see seems to derive from it by duplication to 24 Bouvier 1899). folds per segment. Macroperipatus is charac­ Urinary tubercle is different from M geagi, terized by the quadrangular base of primary where it is overloaded inward and from M dorsal papillae, which are separated by straight perrieri and M. acacioi, because M valerioi has grooves that run parallel with axis of body ; the tubercle, separated from 3 and 4 arcs (Fig. body folds apparently divided by grooves, and 2a,b) while i� the fo rmer three species the tu- accessory papillae usually few and small (peck, * However, the jaws are variable lo 1975). With the exception of accessory papillae, in M. torcuatus, !he extent �at one individual may diffe r in left M valerioi fits in Macroperipatus. Bouvier and righ t jaw structure (H. Ruhberg 1985: pers. (I899b) mentions the high variability in size of ·comm). MORERA & LEON: Ma croperipatus valerioi sp. 279

which has close morphological afinities with M. geagi .

RESUMEN

Se describe Macroperipatus valerioi, una nueva especie de Río Damitas, 16 Km norte de Puerto Quepos, en la costa del Pacífico de Cos­ ta Rica. M. valerioi, sp. n., está relacionado con M. geayi (Bouvier) de Sur América, del cual di­ fiere por poseer un color pardo uniforme en el dorso; por la curvatura de la delgada cuarta al­ mohadilla pedal de los pares cuarto y quinto de lobopodios (ésta rodea al tubérculo nefridial que permanece totalmente libre y recargado al lado posterior); por la presencia de dos dientes accesorios, el segundo con menor desarrollo que el primero, y por tener de 12 a 13 dentícu­ los en la sierra de la mandlbula interna. El tipo es una hembra de 34 pares de patas.

ACKNOWLEDGMENTS

We thank H. Levi, E. Orozco, N.I. Platnick, D. Quintero, H. Ruhberg y A. Wille who kindly allowed use of specimens from their museums. M. Rojas prepared the photographs.

REFERENCES

Amaral, A.D.; S. Campiglia & R. Lavallard. 1980. Study of the hemolymph in Peripatus acacioi (Ony­ ehophora): Plasma total and fraetionated protein levels. C.R. Herb Seanees Acad. Sei. Ser. D. Se. Nat. 290:1I81-1184. Fig. 1. DP: main tooth, DA. aeeessory teeth, DIA: diasterma, D: denticles. (a) Ma croperipatus valerioi: Arnett, R.H. 1961. The Onyehophora of Jamaica. Ent. outer blade (40x) (b) labrum (52x) (e) inner blade, News, 72:213-220. (40x); Macroperipatus torcuatus, (d) outer blade, (e) inner blade, (40x), (Bouvier, 1905); Macroperipatus Bouvier, E.L. 1899. Comptes Rendus Aead. Sei. Paris, outer blade, (g) inner blade, (no scale), guianesis: (f) vo\. 128, p. 1345. (Evans, 1903); Ma croperipatusacacioi: (h) outer blade, Bouvier, E. L. 1899b. Contributions a I'Histoire des (i) inner blade, (bar . 100 ¡.tm) (Mareus & Mareus, 1955); Macroperipatus perrieri (j) outer blade, (k) Peripates Amerieains. Ann. Soe. ent. Fr., 68: 386- inner blade, (l06x), (Bouvier, 1899). 450. Bouvicr, E. L. 1905 . Monographie des Onychophores l. bercles are united (Fig. 2e,g) or free but closely Ann. Sci. Nat. Zoo\. (ser. 9) 2:1-383. surrounded (Fig. 2c,d). On August 10, 1966, S. Peck collected at Brues, C.T. 1911. A new speeies of Peripatus from Grenada, with o bservations on other speeies of the Rincón de Osa, Costa Rica (under a log, 50 m genus. �ull. Mus. Comp. Zool, 54:305-318. 1925. altitude, Tropical wet fo rest, Holdridge, 1967), Brues, C. T. 1925. Notes on Neotropieal Onyehopho­ an onychophoran which he determined as M. ra. PSY he 32: 159-165. geagi (originalIy described from French Guiana). l Campiglia, S.S. 1976. The blood of Peripatus acacioi However, we suspect that the low vagility ofthe Mareus and Mareus (Onyehophora): III. The ionie group precludes genetical flux in large ranges, eompo�ition of the hemolymph. Comp. Biochem and Peck's specimen may rather be M. valeripi, Physiol a Comp. Physiol 54: 129- 133. 280 REVISTA DE BIOLOGIA TROPICAL

Fig. 2. Position 01' nefridial tubercle in 4 and 5 pairs of lobopods. (Iower view). UN: nail, PP: pedal papilla, PI: "foot", PD: dermal papilla, AP: (pedal) ereeping pad, TN: neCridial tubercle. M valerioi: (a) Courth leCt lobopod, (b) fifth right lobopod (barr: 50 Ilm),M. geagi: (e) fifth right lobopod of a Colombian speeirnen, (d) fifth right lobopod 01' the type (from Freneh Guiana), 27.5x, (Bouvier, l 899); Macroperipatus perrieri: (e) fourth left lobopod, (31x), (Bouvier, 1899), Macroperipatus guianensis: (f) fourth lobopod, (no scale) , Evans, 1903);Macroperipatus acacioi: (g) fourth lobopod, (bar : 501lm) , (Mareus & Mareus, 1955).

Campiglia, S. & R. Lavallard. 1973. Contribution a Clark, A.H. 1937. On sorne Onychophores from the la biologie de Peripatus acacioi Mareus et Mareus: West Indies and Central America. Proc. United 11. Variations du poids des animaux en fonetion du Sta tes Nat. Mus. 85 : I -3. sexe et du nombre des lobopodes. Biol. Mar. (Nova Ser.) 30 : 499-5 I 2. Clark, A.H. & J. Zetek. 1946. The Onychophores of Panamá and the Canal Zone. Proc. United States Campiglia, S. & R. Lavallard. 1975. Contribution a Nat. Mus. 96 : 205-213. l' hématologie de Peripatus acacioi Mareus et Marcus (Onychophore): 11. Structure et ultrastrue­ Evans, R. 1909. On Peripatus guianensis (sp. nov) ture des globules péricardiaux. Ann. Sei. Nat. Zool. Quart. Jour. Micr. Sci (N.S.), 47 (2): 145-159. Biol. Anim. 12 Sér 17:93-1 20. Holdridge, L. R. 1967. Life zone ecology. Tropical Campiglia, S. & R. Lavallard. 1982. Water loss in Science Cente�. San José. 206 pp. Peripa tus acacioi (Onyehophora) during experi­ mental inmobilization. Vie Milieu 32: I 7 I -174. Kennel, J. von. 1885. Entwicklengsgeschichte von Peripatus Edwardsii Blanchard und Peripatus C1ark, A.H. 1913a. A revision of the American species torcuatus sp., Abh. Zool. Inst. Wfuzburg 7: 95- 1n . of Peripatus. Proc. Biol. Soco Wash. 26 : 15-20. 229; 1886• !bid. 8: 1-93. 3b. Notes on American species of Clark. A.H. 191 Lavallard, R. 1981. Ultrastructural observations on with a list of the known forms. Smith· Peripatus, the segm�ntal organs oC Peripatus acacioi (Ony- sonian Mise. ColI. 60: 1-5. MORERA & LEON: Ma craperipatus va/eriai sp. 281

Hg. 3. Upper view of dorsal teguments. Arrow: middle dorsal lineo (a) Macraperipatus va/eriai (26x); (b) Ma­ craperipatus perrieri (31 x), (Bouvier, 1899); (e) Ma craperipatus guianensis (no seale), (Evans, 1903); (d) Ma ­ craperipatus acaciai (bar : 200 !lm), (Marcus & Marcus, 1955); (e) Ma craperipatus tarquatus (20x), (Bouvier, 1899); (f)Ma craperipatus geayi (24x), (Bouvier, 1899). ehophora: Peripatidae) Ann. Sci. Nat. lool. Bio. Anim. 3:23-62.

Lavallard, R. & S. Campiglia 1973. Contribution a la biologie de Peripatus acaciai Marcus et Marcus: 1. Poureentage des sexes et variations du nom bre des lobopodes dans un éehantillonnage de plusieurs centaines d'individus. Bol. lool. Biol. Mar (Nova Ser.) 30:483-498.

Lavallard R. & S. Campiglia. 1974. Contribution a l'hématologie de Peripatus acaciai Mareus et Mareus (Onyehophore): 1. Strueture et ultrastruc­ ture des hémoeytes. Ann. Sei. Nat. lool. Biol. Anim. 12 Sér. 17:67-92.

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Lavallard, R. & S. Campig1ia: 1975b. Cont. a la Biol. de Peripatus acaciai M. et M.-V. Etude des naissances dans un élevage de laboratoire. lool. Fig. 4. Lateral view of dorsal papillae in adults (no Anz. Jena 195:338-350. scale). Ma craperipatus va/eriai: (a) principal papillae, (b) accessory papillae; Macraperipatus geagi: (e) prin­ Lava1lard R., S. Campiglia, E. Parisi Alvares & cipal papillae, (d) accessory papillae; Macraperipatus C.M.C. Valle. 1975 Contribution to the biology of acaciai: (e) principal papillae, (f) accessory papillae, Peripatus acaciai Mareus and Mareus (Onyeho­ (Marcus & Marcus, 1955); Macraperipatus insularis: phora)t 111. A descriptive study of the habitat. Vie (g) principal papillae, (Arnett, 1966); Ma craperipatus Mi1ieu Ser. C. Biol. Terr. 25:87-118. i. clarki: (h) principal papillae, (Arnett, 1966); Ma -· craperipatus guianensis: (i) principal papillae, (Evans, Mareus, E. & E. Mareus. 1955. A new Peripatus from 1903); Macraperipatus perrieri: U) principal papillae, Minas Gerais, Brazil, Onais Acad. Brazil. Cien., (Bouvier, 1899). 27: 1 89-293. 282 REVISTA DE BIOLOGIATROPICAL

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