B., J. Monge-Nájera & R. Sáenz. 1988. Parturition
in onychophorans: new record and a review. Brenesia 29: 15-20.
in the Costa Rican Epiperipatus hilkae has the following sequence:
the mother showed peristalsis and a swollen gonopore, the terminal
lobopods are raised and the neonate walks out, a drop of whitish
material is expelled by the gonopore and the young crawled to the
mother«s back. Four young measured 1.2-1.8 mm in length and
weighed 0.010-0.017 g (mean 5.9 % of mogher«s weigth). During
the first six weeks young are more lightly colored than adults.
In onychophorans, the parturition sequence is realtively constant
throughout the taxa, requires 10-30 min and its frecuency appears
to be influenced by environmental conditions. The systematic distribution
of modes of reproduction suggests and increasing tendency towards
greater parental investment.
& J. Monge Nájera. 1990. Epiperipatus hilkae, n. sp.
from Costa Rica (Onychophora: Peripatidae). Rev. Biol. Trop. 38(2B):
hilkae, a new species discovered in tropical dry and moist forests
(NW Costa Rica) has the following characteristics: in the fourth
and fifth oncopods, the nephridial tubercle is free from the third
sole, and only partially surrounded by the fourth arc. Each segment
has 12 skin folds (seven folds reach the ventral side), which are
divided only by the midline. The primary papillae are conical with
rouded bases and two to three scale ranks in the apical section.
The external jaw blade has two accessory teeth, the internal with
one accessory tooth and 12 denticles. The diastema is monolobular.
The dorsal part is dark brown with a pattern of hexagons conspicuous
for their light reddish-brown papillae. The new species is closely
related to E. isthmicola and "E. nicaraguensis", also
from Central America. It remains active during the dry season in
isolated moist patches.
J., Z. Barrientos & F. Aguilar. 1993. Behavior of Epiperipatus
biolleyi (Onychophora: Peripatidae) under laboratory conditions.
Rev. Biol. Trop. 41(3): 689-696.
behavior of Epiperipatus biolleyi Bouvier was studied in the laboratory.
In choice tests, bryophyte vegetation and its associated soil were
preferred to grass and its soil. In 87 hr the animals chaged artificial
burrows 2.89 times. They enter burrows mostly by walking forward
and show a tendency to rest facing the entrance. No agressive competition
for burrows was observed. Pairs rest with some body contact about
half the time. Seven resting body postures were identified. They
hide from direct sunlight in 189 s (mean) when placed over moss
and appear to avoid light around 470-600 nm. Walking speed was near
1cm/s. They float and become tergid in freshwater but drown in sea
water. Their adhesive secretion has a bitter taste and dissolves
in less than 3 s in sea water but remains adhesive under freshwater
for at least 20 hr. In nature, animals bear scars and mutilated
oncopods. Death is often preceded by a retraction of the antennae
and expulsion of saliva, adhesive substance, faeces and sometimes
embryos. Ecdysis occurs aproximately every 15 days. At least one
bird (Lturdus grayii) and one snake (Micrurus hemprichii) are known
to prey on other onychophoran species in the wild.
J. & B. Morera. 1994. Morphological and physiological characteristics
of two species of Epiperipatus from Costa Rica (Onychophora: Peripatidae).
Rev. Biol. Trop. 42 (1/2): 181-188.
biolleyi and Epiperipatus isthmicola from Costa Rica were studied
in the laboratory. Morphometric analyses of E. biolleyi show that:
(1) larger animals have more legs and (2) females have 8% more pairs
of legs, and are 24% longer and 50% heavier than males. An estimation
of how alcohol preservation affects body length indicates less shrinkage
in females. Physiological observation on E. isthmicola showed that
(1) in a session these animals can lose a mean of 7.4% of body weight
(BW) by expelling their adhesive secretion, (2) weight loss by water
evaporation was in average 2% of BW/min at a 66% relative humidity,
and (3) larger animals lose water more slowly. An observed parturition
of E. isthmicola had the sequence and timing that are typical of
the family Peripatidae.
J. 1994. Ecological Biogeography in the Phylum Onychophora. Biogeographica
70 (3): 111-123.
ecological biogeography of onychophorans has been the subject of
some qualitative observations, but the results have been incosistent.
Thes paper presents the results of (1) statistical analyses of 172
geographic quadrats for both onychophoran families (worldwide) and
(2) independent graphic analyses of most South African and Australian
species. Both types of analysis produced different results, including
a correlation of occurrence with Pleistocene vegetation, which does
not imply causation. Although altitude and rainfall co-relate with
the distribution of some South African and Australian taxa, each
of the following factors failed to explain by itself the barriers
that limit onychophoran distribution at the global level: mean annual
rainfall and temperature, photosynthesis, and types of climate,
vegetation and biome. The barriers seem to be a product of several
ecological factors which depend on the onychophoran taxon.
J. 1994. Reproductive trends, habitat type and body characteristics
in velvet worms (Onychophora). Rev. Biol. Trop. 42 (3): 611-622.
quantitative analysis of several onychophoran characteristics shows
that in habitats with lower rain levels females reproduce at an
older age, are more fecund and tend to have reproductive diapause
where rain does not exceed a mean of 200 cm/year. These habitat
characteristics are associated with the southern family Peripatopsidae.
Sex ratio and parental insvestment per yong are not correlated with
general environmental conditions. A comparison of 72 species showed
that larger species are often more variable in morphometry, but
species with the longest females do not always have the longest
males. Larger Peripatus acacioi females (Peripatidae: Brazil) produce
more and heavier offspring. Intrapopulation morphology was studied
in 12 peripatid species for which samples of between 11 and 798
individuals were available. In general, within populations the females
are more variable than males in length and weight, but similarly
variable in the number of legs. The number of legs has a low variability
(1.73-2.45%), length is intermediate (22.4-25.3%) and weight is
very variable (49.41-75.17%). When sexes are compared within a population,
females can have 1.4-8.9% more leg pairs, and be 47-63% heavier
and 26% longer than males.
J. 1995. Phylogeny, biogeography and reproductive trends in the
Onychophora. Zool. J. Linn. Soc. (London) 114: 21-60.
cladistic analysis places the Onychophora between Polychaeta and
Arthropoda. The 'Uniramia' concept is not supported. No justification
was found for either onychophoran family to be considered ancestral.
A cladogram of fossil genera indicates the common ancestor to have
long oncopods, armoured plates and an annulated body. Later forms
show adaptations to life in reduced spaces. Physiological data suggest
that the Onychophora became adapted to land via the littoral zone,
before the Late Ordovician. Adhesive glands evolved for defense
on land. Peripatopsidae and Peripatidae were distinct by the late
Triassic. The occurrence of onychophorans probably dates from post-Early
Cretaceous in Chile, the southern half of Southeast Asia, Mesoamerica
and the Caribbean. After the Early Cretaceous, the peripatids of
tropical Africa lost terrestrial contact with those of South America.
A new biogeographic technique, formalized here under the name retrovicariance,
indicates that the Peripatidae of Equatorial Africa and the Neotropics
are sister-groups. Typical inbreeding adaptation found in some onychophorans
include: female-biased sex ratios; gregarious development; relatively
constant time of development and number of offspring in each clutch;
male polygamy and shorter life span; frequent sibmating in the microhatibat
of development, and sperm storage by females, so that a single insemination
fertilizes all ova.
J. & José P. Alfaro. 1995. Geographic variation of habitats
in Costa Rican velvet worms (Onychophora: Peripatidae). Biogeographica
71 (3): 97-108.
characteristics were compared for 20 onychophoran localities in
Costa Rica, from the seasonally dry western Pacific forest to the
rainforests of the Caribbean. Everywhere, rainfall is more variable
than temperature and than relative atmospheric humidity. A microhabitat
study of Epiperipatus biolleyi Bouvier, 1902 was done for comparison
with the only other species for which equivalent data are available,
the Brazilian Peripatus acacioi. The Costa Rican species was found
(1) in sandy, not clay rich soil, (2) closer to the surface and
(3) in burrows whose temperature es more similar to the external
air temperature. For both species the soil humidity (mean 35 %)
and acidity (pH=5.2-6.2) were similar. The maximum E. biolleyi population
density was 0.25 individuals/m2. No clearcut trends in associated
flora and fauna were found. In the laboratory, the animals preferred
rotten to non-rotten wood, and water-soaked soil to oven-dried soil,
during periods of inactivity.
J. & W. Lourenco. 1995. Biogeographic implications of evolutionary
trends in onychophorans and scorpions. Biogeographica 71(4): 179-185.
comparison with another group of terrestrial predatory invertebrates,
the Order Scorpiones, suggests that the lack of adaptations to dry
microenvironments has been the central limitation to further biogeographic
radiation in the phylum Onychophora.
J. 1996. Jurassic-Pliocene biogeography: testing a model with velvet
worm (Onychophora) vicariance. Rev. Biol. Trop. 44(1): 159-175.
paleomaps of global continental vegetation from the Jurassic through
the Pliocene were prepared (based on the literature) and used to
define an area cladogram. Bouvier«s (early XX century) natural
classification of velvet worms (phylum Onychophora) was used to
define an independent taxonomic/geographic cladogram of onychophorans
for all regions where they are found today. Both cladograms show
the same sequence of geographic vicariance. Thus, the paleogeographic
model is supported by the taxonomic evidence. The paper includes
a color atlas.
J. , Z. Barrientos & F. Aguilar. 1996. Experimental Behaviour
of a Tropic Invertebrate: Epiperopatus biollegi (Onychophora: Periapatidae).
In J.J. Geoffroy, J.P. Muries & m. Nguyen Duy-Jacquemin (eds).
Acta Myriapodologica. Mem. Mus. nath. Hist. nat. 169: 493-494.
basic behaviour of Epiperipatus biolleyi Bouvier 1906 was studied
experimentally in the laboratory with specimens collected in the
central plateau of Costa Rica (Coronado area of San José
province). The animal avoids light of a wave length between 470
and 600 nm. Perhaps it lacks the ability to detect light in the
infrared and ultraviolet light range, suggesting that the wider
visual range of insects- for instance- was acquired later in arthropodan
and A.C.C. Wilson, 1999. Microsatellite markers for the onychophoran
School of Biological
Sciences, Macquarie University, NSW 2109, Australia
species, local endemism, mating system, peripatus. email@example.com
We are studing
Euperipatoides rowelli (Reid 1996) as a model of population processes
at increasing spatial scale from local patterns of mating through
to regional phylogeography. This onychophoran is cornmon in rotting
logs in forests of southeastern New South Wales (NSW), Australia
(Reid 1996). Allozymes identify regional cryptic species, but are
unusually invariant at finer scales (Tait et al. 1995), and therefore
we cloned microsatellites which could be expected to be more variable.
conditions, only loci P6 and P23 gave scoreable polymorphic patterns
of the expected size (Table 1). (These loci work well in duplex
if the primer concentrations of P6 are doubled.) Primers for P17
gave poor aniplification of the (CA)14 170 bp target, but yielded
two other polymorphic loci: 'P17 low' and 'P17free' (there is no
linkage disequilibrium between these, P > 0.05 in all populations).
Shorter P17 primers (Table 1) give better results for P17low. P18
could not be made to amplify until we added 0.01% final Tween 20
and Nonidet P40 detergents, with which the locus is scoreable. Consistency
with Mendelian inheritance of the five loci has been confirmed in
mother-offspring and population samples.
Some loci do
not amplify at all in sorne populations (Table 1), probably because
E. rowelli is a species complex: adjacent populations can differ
by 20% (or more in mtDNA COI sequence (P. Sunnucks, unpublished).
P6 and P23 have nonamplifying alleles, including in the population
from which they were cloned.
and F. Sunnucks, 1999. Molecular anatomy of an onychophoran: companmentalized
sperm storage and heterogeneous patemity. Molecular Ecology, 8,
School of Biological
Sciences, Macquarie University, Sydney 2109 NSW Australia
rowelli, female control, multiple paternity, Onychophora, spermathecae
(peripatus or velvet worms) show extraordinarily high local endemism,
and cryptic species are common. As part of a programme addressing
issues of endemicity at hierarchical spatial scales, we investigated
reproduction in Euredratoides rowelli (Onychophora: Feripatopsidae)
using microsatellite analysis. This species is ovoviviparous, and
females have up to 70 embryos in their uten simultaneously Batches
of undeveloped and well-developed embryos may be present in the
uten of a female. Paired ovaries lead via a common oviduct into
paired uteri, each of which has a spermatheca (sperm storage organ).
Insemination in E. rowelli is dermal-haemocoelic: spermatophores
are placed on the skin of the female, the body wall is breeched,
and sperm are released into the haemocoel through which they migrate
to the spermathecae. There is no obvious mechanism to prevent sperm
mixing, yet microsatellite analysis indicated that offspring in
a female's paired reproductive tracts can be sired by different
males, and that the paired spermathecae can contain sperm from different
males. More than 70% of females had broods with multiple paternity.
The data are consistent with the potential for female postcopulatory
influence over fertilizations: in particular, compadmentalization
of sperm from different males into different spermathecae. Female
control of fertilizations could lead to benefits including increased
diversity of offspring, minimization of maternal-paternal genetic
incompatibility, and influence on offspring genotvpes. Multiple
mating alone may increase Ihe genetic diversity of offspring: this
could be of importance in E. rowelli, which has very small genetic
neighbourhoods and low genetic marker diversity.
J. & R. Ramírez. 1996. El phylum Onychophora en el Perú.
V RC ICBAR (Perú) March: 74
following new records are report for Peruvian tropical forest: Oroperipatus
weyrauchi at Parque nacional Yanachaga-Chemillén, Pasco (one
specimen); Oroperipatus bluntschlii from Río Lagartococha,
Loreto (three specimens) and Oroperipatus quitensis, Cordillera
del Cóndor, Amazonas (one specimen). They are deposited in
the Entomological Collection, Museo de Historia Natural, Universidad
Mayor de San Macos, Perú.
Jr. 2000. Fossil onychophorans from Dominican and Baltic amber:
Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis
balticus n.g., n.sp. (Succinipatopsidae n.fam.) with a proposed
classification of the subphylum Onychophora
dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis
balticus n.gen., n.sp. (Succinipatopsidae n.fam.) (Lobopodia: Onychophora),
the first Tertiary fossils of the Lobopodia, are described from
Dominican and Baltic amber; respectively. Both families are characterized
by the presence of simple legs lacking foot portions with claws
and pads. Tertiapatidae is further characterized by soluble body
pigments and oral papillae shorter than the legs. Succinipatopsidae
is characterized by non-soluble body pigments and oral papillae
longer than the legs. Nomenclatural changes include the erection
of the class Udeonychophora n.nom. for terrestrial onychophorans
with a ventral mouth, the order Ontonychophora n.nom. for extant
onychophorans possessing legs with a differentiated "foot"
portion, and the family Helenodoridae n.nom. for the genus Helenodora
from the Carboniferous. The biogeographical significance of these
fossils and their phylogenetic relationship with previously described
onychophorans are discussed.
words: Euonychophora, fossil velvet worms