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Rediscovery of Boulengerula fischeri, with notes on its morphology andhabitatG. John Measeyab; Harald H. Hinkelc; Bonny Dumbod; Eberhard Fischerda Applied Biodiversity Research, Kirstenbosch Research Centre, South African National BiodiversityInstitute, Claremont, South Africa b Department of Biodiversity and Conservation Biology, Universityof the Western Cape, Bellville, South Africa c The World Bank Country Office Kigali, Kigali, Rwanda d

Institute for Integrated Natural Sciences, Department of Biology, University of Koblenz-Landau,Koblenz, Germany

Online publication date: 15 March 2011

To cite this Article Measey, G. John , Hinkel, Harald H. , Dumbo, Bonny and Fischer, Eberhard(2011) 'Rediscovery ofBoulengerula fischeri, with notes on its morphology and habitat', African Journal of Herpetology, 60: 1, 47 59

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Original article

Rediscovery of Boulengerula fischeri, with notes on its

morphology and habitat

G. JOHN MEASEY1,2*, HARALD H. HINKEL3, BONNY DUMBO4

& EBERHARD FISCHER4

1Applied Biodiversity Research, Kirstenbosch Research Centre, South African National Biodiversity Institute,

Claremont, South Africa; 2Department of Biodiversity and Conservation Biology, University of the Western

Cape, Bellville, South Africa; 3The World Bank Country Office Kigali, Kigali, Rwanda; 4Institute for

Integrated Natural Sciences, Department of Biology, University of Koblenz-Landau, Koblenz, Germany

Abstract.*

Caecilians remain an enigmatic component of the tropical terrestrial fauna duein part to their cryptic life-history but also to a lack of directed effort towards their study.

Many species are known from single specimens associated with poor collection data, which

has not facilitated re-collection. Consequently, nearly two thirds of species are currently

considered Data Deficient by the IUCN. Boulengerula fischeri was described from a single

specimen deposited onto a car by an unknown collector in 1987, and has defied attempts at

relocation for nearly 20 years. Herein we describe a collection of 20 individuals from

agricultural land immediately outside Cyamudongo Forest in southwestern Rwanda,

around 3 km from the type locality. Adults were found to attain a remarkable maximum

length of nearly 400 mm despite being only 4.5 mm wide (maximum elongation index of

102). The range of morphometric and meristic measurements taken include those given for

the type specimen, featuring mean annular and vertebral counts of 199.4 and 191.8,

respectively. The estimated density of B. fischeriwas found to be 50 animals h(1 km(1 and

is comparable with other species of Boulengerula. Despite our new collection records, we

recommend that this species remains Data Deficient until its range can be better described.

Key words.*Gymnophiona, Central Africa, Rwanda, soil megafauna, body elongation

Caecilians have been described as rare enigmatic components of the tropical terrestrial

ecosystem (e.g. Gundappa et al. 1981; Bhatta 1997). Such views are easily justified

when considering the general paucity of collections of these animals (e.g. Taylor 1968).

However, recent targeted collections have shown that as a generality caecilians cannot

be considered rare (e.g. Measey 2004), suggesting that applied searches of areas in

African Journal of Herpetology,

Vol. 60, No. 1, April 2011, 4759

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seven currently recognised species (Muller et al. 2005). Some are known from good

collections with well established ecology and natural history: B. boulengeri

(e.g. Gower et al. 2004; Measey 2004; Jones et al. 2006; Measey & Barot 2006)

and B. taitanus (e.g. Hebrard et al. 1992; Gaborieau & Measey 2004; Measey 2004;

Malonza & Measey 2005). Others are known from few specimens or from sporadic

collections: B. chamgamwensis (see Malonza & Muller 2004) and B. uluguruensis (see

Measey et al. 2006). Lastly, two species are known only from the holotype: a single

specimen of both B. denhardti and B. fischeri (Nussbaum & Hinkel 1994; Wilkinson

et al. 2004).

The taxonomic status of some Boulengerula species remains unclear. For example,Nussbaum & Hinkel (1994) described the distribution of B. boulengeri from the

Usambara and Magrotto Mountains, northeastern Tanzania, and Malonza & Muller

(2004) recognised B. boulengeri only from the East Usambaras, while Channing &

Howell (2005) suggested an additional species from the West Usambaras. Malonza &

Muller (2004) questioned whether B. chamgamwensis from Malawi are the same as

those found in coastal Kenya (see Nussbaum & Hinkel 1994). At the generic level,

the proposed synonomy Boulengerula and Afrocaecilia (see Nussbaum & Hinkel1994) has been challenged as premature (Wilkinson et al. 2003). Certainly, for both

morphological and molecular phylogenetic work, more specimens are needed. In

this respect, central Africa is perhaps one of the most interesting areas as it has

already produced two caecilian specimens: an unverified record of Schistometopum

tentatively ascribed to S. thomense (see Nussbaum & Pfrender 1998; Measey et al.

2007), and the monotypic B. fischeri.

Boulengerula fischeri was formally described by Nussbaum & Hinkel (1994: 756)from a single individual, an immature female collected 0630 h. 29 April 1987 at

Foret de Cyamudongo near Foret de Nyungwe, 17002000 m, Cyangugu Prefecture,

Rwanda by native collectors for E. Fischer. No paratypes. However, the exact

circumstances around which this specimen was found reveal some uncertainty as to

its provenance. In April 1987, one of us (EF) had asked local people in the area

adjacent to Cyamudongo Forest to collect earthworms for a colleague who wished to

make a taxonomic study. A great many earthworms were brought on the 27 and

28 April 1987, so that collectors were asked to stop providing them. However, on

returning to the car which was parked in a clearing inside Cyamudongo Forest on

29 April, several earthworms were found draped across the bonnet. It was concluded

(by EF and HHH and as reported by Nussbaum & Hinkel (1994)) that the

earthworms had been brought in response with the earlier request, but that the

48 MEASEY ET AL.**Rediscovery of Boulengerula fischeri


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to collect earthworms, but no more specimens of B. fischeri were found, although a

great many earthworms were offered. While collection of earthworms seemed the

most likely explanation for the appearance ofB. fischerion the car bonnet on 29 April

1987, the continued passing of years without another specimen raised suspicions

about possible alternative hypotheses for the appearance of the holotype. Could it

have been dropped by a passing bird? Or as one colleague earnestly suggested, perhaps

this caecilian is specialised for living in arboreal moss balls and had fallen from a tree

overhead; that arboreal moss balls do not occur in Cyamudongo Forest did not deter

the proponent of this hypothesis! Although the exact details for the original collection

of the holotype of B. fischeri will probably always remain unknown, it would clearlybe possible to find out a great deal more about this species from new collections.

Nussbaum & Hinkel (1994) remark that B. fischeri was one of the most unusual

Boulengerula due to the combination of its very high annular and vertebral count

(186 and 194, respectively) and a very slender body. They also noted that nothing is

known about the natural history of this species. Further, in the context of

conservation, Gower & Wilkinson (2005) called for priority to be made for assessing

caecilian species which had not been reported for many years, especially in the case ofmonotypic taxa. Our objective was to discover exactly where B. fischeri occurs and

increase scientific knowledge of the only caecilian ever to be described from central

Africa.

MATERIALS AND METHODS

Study Site

A detailed description of the flora and fauna of Rwanda including the study site and

surrounds is given by Fischer & Hinkel (1992). Below we summarise information

pertinent to this study.

Rwanda is a small country in central Africa between 18 04? and 28 51? latitude

South and 288 53? and 308 53? longitude East. Despite its small size (26 338 km2),

Rwanda occurs at one of the most important watersheds on the African continent.The source of the White Nile is found in central Rwanda and from there flows north,

while rain in the south is divided between the Congo basin westwards and the

Albertine Rift to the east. The climate fluctuates greatly with altitude from a mean

temperature of 248 C at 1 000 m asl to 08 C at the summit of Karisimbi (4 507 m asl),

h hi h f h i h j l f h Vi fi ld i h R d

AFRICAN JOURNAL OF HERPETOLOGY 60(1) 2011 49


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Cyamudongo Forest has a rich endemic flora (Fischer & Hinkel 1992). For a detailed

description of Nyungwe National Park see Fischer & Killmann (2008).

Cyamudongo Forest, today also part of the National Park, is surrounded by

low-intensity farming, typically comprising of bananas (Musa sp.), avocado (Persea

americana Mill.), sugar cane (Saccharum officinarum L.), yams (Dioscorea sp.),

tomatoes (Lycopersicon esculentum Mill.), beans (Phaseolus vulgaris L.), cassava

(Manihot esculenta Crantz) and onions (Allium sp.) interspersed with single trees of

the once larger Cyamudongo Forest (e.g. Entandrophragma excelsum). The only

nearby intensive agriculture is a tea plantation (Camellia sinensis) 3 km to the north.

Searching for Boulengerula fischeri

Although the locality where the holotype of B. fischeriwas found was in a clearing in

Cyamudongo Forest, we suspected that the animal had been unearthed in an

adjacent agricultural area, as is known for most of the genus (see Malonza & Muller

2004; Measey 2004; Malonza & Measey 2005; Measey et al. 2006), and thereforeconcentrated our limited searching time to the forest near the clearing as well as

agricultural areas immediately around the forest. We encouraged the help of local

people by showing them a picture ofB. fischeri(see Fischer & Hinkel 1992: 115) and

describing the difference from earthworms and snakes: presence of a hard skull,

presence of mouth, does not change length or girth, and absence of scales.

In March 2006, we dug with hoes (locally known as amazuka) in appropriate

microhabitats: under rotting logs, in soft soil, in any deep accumulations of leaf litter

(such as within the buttresses of trees or under banana plants), or at any particularly

wet area (in and around a seepage, or next to a stream); we paid particular attention

to digging around the base of banana plants (see Measey 2004).

Measurements at collection sites included soil texture (Dubbin 2001), soil

temperature using a temperature probe set 10 cm into the soil, and soil pH made

by mixing approximately 1:2 volume with stream or well water (the pH of which was

measured independently to avoid extreme acid or alkaline water). Temperature and

pH were measured with an Acorn pH 6 (Forestry Suppliers, USA). Three thermo-

button temperature loggers (Proges Plus, France) were placed at the site of the first

caecilian found and left for 24 hours: 30 cm deep in the soil, 5 cm deep under the soil

surface, and 1 m above soil surface on a shaded banana stem. Shade was estimated

with regard to the total area searched or surveyed over a site, and attention was also


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digital images of radiographs. Specimens are deposited in Zoologisches Forschungs-

museum Alexander Koenig (ZFMK 91849-55).

RESULTS

We investigated a total of six sites in an area of 5.5 km2 in and around Cyamudongo

Forest over two days in March 2006 (Table 1a, Fig. 1). Boulengerula fischeri were

only found at one site, despite showing images and live animals to local people. Sites

explored were natural forest or contained similar crops (see above).Most local people did not recognise the picture shown as being anything other

than an earthworm. When we explained that it was not an earthworm, but gave a

description (see above), most people claimed not to know the animal and lost interest

in helping. The only exception to this was one woman from Nyaruhondo village who

claimed to have seen a similar animal that morning moving over the soil surface

amongst her onions, this claim was later verified when she was shown a living

specimen of B. fischeri. Other people from the same village who were shown liveanimals appeared to have no problem recognising and finding animals. However,

people from Muhange who were shown live animals and asked to collect more from

their agricultural plots returned only earthworms. We did not find any B. fischeri

within the forest around the type locality (itself a clearing, see above). The following

results refer to areas where B. fischeri was found:

Soil Measurements

Soil temperatures measured from microhabitats where B. fischeri was found appear

to agree with soil temperature data retrieved from thermo-buttons. The 24 h

temperature profiles indicate that for all of the night hours, including dawn and

dusk, the air temperature is below that of soil at 30 cm. Minimum air temperatures

were 16.58 C from 01h30 to 06h15, while maximum air temperatures of 228 C were

found around midday, giving a range of 5.58 C. Just below the soil surface (5 cm) therange of temperatures recorded were 17.5 to 19.58 C, following a lag of around

1 hour from air temperatures. At a depth of 30 cm the temperature remained

constant at 19.58 C. All soils where B. fischeri were found had a clay loam texture

with a pH from 7.44 to 7.63.


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Table 1. (a) Sites searched for B. fischeri around and at the type locality in Cyamudongo forest (see Fig. 1). (b) Details of the sublocalities where

caecilians were collected at Nyaruhondo village. Means (9standard error) are given for temperatures, depths and penetration for each caecilian found

(n020).

(a)

Site DateArea searched

(m2)Time spent (person

hours) Shade % B. fischeri AltitudeDistance to type(km)'direction

Type locality 24/03 250 1 100 0 1994 0

Muhange 24/03 2500 3 60 0 2060 1.3 N

Muhange West 25/03 400 2 80 0 1920 2.2 NW

Muhange East 25/03 600 2 60 0 2000 1.1 NNE

Nkungu 25/03 400 1 80 0 1800 2.9 NE

Nyaruhondo 25/03 1100 3 70 5 1825 2.9 ENE

(b)

Site no. Soil tempDepth range

(cm) Slope facing Shade % B. fischeri AltitudeMean penetration

(kg.cm(2)

Nyaruhondo Valley 18.4 015 South 40 7 1737 3.83

Village 2 18.9 823 ENEast 70 7 1778 5.56

Bosco 19.3 515 WNWest 50 3 1813 3.7Village 19.1 1833 North 70 3 1825 4.36

Mean9SE 18.790.30 11.794.30 4.790.32

52

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ETAL.**RediscoveryofBoulengerulafischeri

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morning of 26 March, a further 12 animals were unearthed by local people at four

sub-localities totalling some 7500 m2 all within a 200 m radius of the eight animals

caught on 25 March.

The holotype of B. fischeri is an immature female measuring some 191 mm in

total length (Nussbaum & Hinkel 1994). The results of our collection show that this

species reaches nearly 400 mm in total length (measured live, range 173397; mean

269.7 mm; Table 2; Online Supplementary Material), making it by far the longest

recorded of any Boulengerula species. However, adults are also extremely attenuate

with a maximum mass of only 4.8 g. On dissection, two females were found to have

Figure 1. Locality of collections of Boulengerula fischeri (circles) in south-westernmost

Rwanda with dark lines indicating national borders and the southern tip of Lake Kivu

(northwest corner). Shaded areas are Cyamudongo Forest and the larger Nyungwe Forest.

Star indicates type locality given in Nussbaum & Hinkel (2004) and cross at position of

Muhange village where B. fischeri could not be found.


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Table 2. Morphometric and meristic measurements of a sub-sample of Boulengerula fischeri

collected from Nyaruhondo village, Rwanda. For each measurement means (mm) aregiven9standard error followed by the range. Sexes are given for mature adults and other

individuals are collectively termed immature. Total length (mm) and mass (g) were measured

in the field, and these data are included in addition to all other measurements made on

preserved specimens, following Wilkinson et al. (2004). Type measurements come from

Nussbaum & Hinkel (1994).

Type Female Females (n03) Males (n03) Immature (n08)

TL (live) 342.6943.34 347.6920.28 213.1098.46

257397 317386 173247

Mass 3.3390.81 3.6390.59 1.3590.19

1.74.2 2.94.8 0.82.4

Preserved Total Length 191 314.0938.59 328.70920.58 203.0098.43

240370 295366 164236

Total annuli 186 196.390.33 190.7090.67 190.6091.24

196.00197.00 190192 185195

Total vertebrae 194 201.7093.33 199.0091.73 198.6091.56

195.00205.00 196202 192205

Distance between tip of 3.8090.34 4.3090.08 3.2090.14

snout and jaw angle 3.164.28 4.234.47 2.803.92

Distance between tip of 2.9090.30 3.1090.02 2.6090.08

lower jaw and jaw angle 2.333.26 3.033.09 2.332.90

Head width at level of jaw 2.8 2.9090.24 3.0090.07 2.3090.05

angles 2.373.16 2.943.16 2.072.55

Head width at 2.9090.31 3.1090.06 2.4090.06

anteriormost nuchal

collar groove

2.313.36 3.013.20 2.102.63

Distance between tip of 5.6 4.9090.33 5.1090.09 4.0090.09

snout and anteriormost

nuchal collar groove

4.225.21 4.945.26 3.754.57

Length of first nuchal 1.7090.07 1.8090.07 1.4090.05

collar, measuredlaterally 1.61

1.84 1.75

1.98 1.20

1.62

Length of second nuchal 2.0090.06 2.0090.10 1.5090.04

collar, measured

laterally

1.932.14 1.882.21 1.371.75

Distance between 1.0090.10 1.1090.06 1.0090.03

i i f 0 80 1 10 1 00 1 22 0 87 1 12


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DISCUSSION

In recent years, several caecilians have been rediscovered following long periods

without collection (e.g. Malonza & Muller 2004; Measey et al. 2006; GJM

unpublished data). It should be stressed that in the majority of cases, the caecilians

in question have been present, and even abundant, throughout the intervening period

but have gone uncollected due to a lack of interested collectors more than any other

reason.

It is notable that Boulengerula fischeri, like other members of this genus, wasfound in small-scale banana plantations (Malonza & Muller 2004; Measey 2004;

Malonza & Measey 2005; Muller et al. 2005; Measey et al. 2006). This type of

microhabitat seems amenable to these and many other caecilians (e.g. Oommen et al.

2000; Deletre & Measey 2004; Measey & Van Dongen 2006), perhaps due to the high

Table 2 (Continued)

Type Female Females (n03) Males (n03) Immature (n08)Distance between naris 1.1090.11 1.0090.08 0.8090.04

and margin of upper lip 0.931.28 0.881.14 0.720.98

Distance between tentacle 1.8090.22 1.8090.08 1.5090.03

and jaw angle 1.412.18 1.651.91 1.401.67


and tip of snout 1.452.12 1.792.09 1.381.66


and margin of upper lip 0.300.48 0.400.46 0.180.33Width at midbody 3 3.8090.43 3.5090.04 2.7090.11

3.014.50 3.423.57 2.363.28

Width at anterior margin 3.5090.51 3.6090.06 2.5090.12

of terminal shield 2.524.22 3.503.69 2.213.32

Length of terminal shield, 3.4090.04 3.2090.18 2.4090.13

measured laterally 3.343.48 2.813.41 2.013.16

Circumference at 11.1090.93 11.6090.84 8.7090.39

midbody 9.3012.40 10.213.1 7.5010.8

Circumference 5 annuli 10.0090.61 10.6090.55 8.0090.24

anterior to terminal

shield

8.8010.80 13.111.5 7.39.4


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very close by in Nyaruhondo (Table 1). We also consider that more time spent

searching within Cyamudongo forest would yield B. fischeri specimens.

We found that there was considerable difference in attitude towards B. fischeriand the collection of any subterranean vertebrates and invertebrates between the

people of Muhange, and the indigenous population, formerly known as Abatwa, of

Nyaruhondo and surrounds. The people of Nyaruhondo are thought to be

indigenous to the Congolian forests with considerable hunting and collecting

experience. They examined both photographs and specimens intently before

attempting any search. The villagers did not have a local name for B. fischeri, but

recognised it as being different from earthworms and snakes. Although they knewthat the caecilian was harmless, many people showed great reluctance to handle live

animals. In their subsequent searches, we were not presented with any earthworms.

In contrast, despite being shown live specimens together with simple diagnostic

features (see above), people from Muhange simply collected earthworms and were

surprised that this is not what was wanted. We consider that it is most likely that the

holotype of B. fischeri was collected by someone in the Nyaruhondo area, who,

during the late 1980s, frequented Cyamudongo Forest (HH & EF pers. obs.). Thisexplanation is consistent with our results in terms of distribution; we only found

B. fischeriin Nyaruhondo village. We feel that our own excavations around Muhange

would likely have revealed B. fischeri had they been present at similar depths and

densities and that they would certainly have been recovered after the annual requests

for earthworms in the same area (see above). However, the exact locality where the

holotype was unearthed remains unknown.

Nussbaum & Hinkel (1994) placed B. fischeriwithin Boulengerula following their

revised diagnosis of the genus. However, they note that the high number of annuli

and vertebrae separate B. fischeri from all other Boulengerula. Perhaps most

surprisingly, B. fischeri attains a maximum length of nearly 400 mm while

maintaining an extremely slender body of less than 5 mm in diameter, providing a

body elongation index of up to 102 (mean 80.8). This remarkable elongate form gives

the live animals the appearance of live pink spaghetti (Figure 2). Herrel & Measey

(2010) found that live animals from the same collection locality were able to perform

internal concertina locomotion to move through narrow tunnels and while burrowingin soil. It is interesting to note that this species was observed moving over the soil

surface, presumably using lateral undulation locomotion. Frequency of surface

movements is thought to be correlated with pigmentation in caecilians (Wollenberg &

Measey 2009). Mean soil penetration was 3.7 to 5.5 kg cm(2 which is relatively high


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cannot currently be made. We also suggest that close attention be paid to possibleupper and lower altitudinal limitations of this species. The nearby mountains in

Democratic Republic of Congo should also be investigated for caecilians, and in

particular to verify whether Schistometopum occurs there (see Nussbaum & Pfrender

1998).

Sub-Saharan Africa represents great potential for collections of many caecilians

known from only a few specimens, extending ranges as well as the discovery of new

species. Nussbaum & Hinkel (1994) give a list of African regions where caecilianshave yet to be collected: northern Malawi, northern Mozambique, northeastern

Zambia and southern Tanzania. They also stressed that searches should not

necessarily be confined to forested areas, but to agricultural areas which may have

been formerly forested, or even other moist habitats. The other area emphasised by

N b & Hi k l (1994) h C h f i l Af i d

Figure 2. An adult female Boulengerula fischeri in life (Total Length0397 mm; Body

Width04.6 mm) showing the extreme elongate nature of this caeciliid caecilian.


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for logistical support. Two referees provided excellent critical comments of a previous

draft. We are grateful to the Office Rwandais du Tourisme et des Parcs Nationaux

(ORTPN) for collection and export permits. Finally, this study would not have been

possible without the support of the people from Nyaruhondo village.

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Received: 2 August 2010; Final acceptance: 16 February 2011


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Rediscovery of Boulengerula fischeri, with notes on its

morphology and habitat

G. JOHN MEASEY1,2, HARALD H. HINKEL3, BONNY DUMBO4 &

EBERHARD FISCHER4

1Applied Biodiversity Research, Kirstenbosch Research Centre, South African National Biodiversity Institute,

Claremont, South Africa; 2Department of Biodiversity and Conservation Biology, University of the Western

Cape, Bellville, South Africa; 3The World Bank Country Office Kigali, Kigali, Rwanda; 4Institute for

Integrated Natural Sciences, Department of Biology, University of Koblenz-Landau, Koblenz, Germany

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Supplementary information for Rediscovery of Boulengerula fischeri, with notes on its morphology and habitat

All measurements are in mm. Sex is indicated for each specimen: m0

male; f0

female; j0

immature

Field Tag JM02573 JM02555 JM02560 JM02571 JM02581 JM02576 JM02561 JM02578 JM02575 JM02568 JM02577 JM02556 JM02570 JM02569

TL (live) 386 397 317 374 257 215 202 189 223 227 173 229 247 340

Mass 4.8 4.1 2.9 4.2 1.7 1.7 1 0.9 1.6 1.2 0.8 1.2 2.4 3.2

Sex m f m f f j j j j j j j j m

Preserved TL 366 370 295 332 240 212 192 176 219 212 164 213 236 325

Total annuli 190 196 190 196 197 188 195 190 193 185 189 195 190 192

Total vertebrae 202 205 196 195 205 205 202 195 195 200 198 202 192 199

Distance between tip

of snout and jaw

angle

4.23 4.28 4.26 4.05 3.16 3.17 2.97 2.81 3.12 3.92 2.80 3.60 3.44 4.47

Distance between tip

of lower jaw and

jaw angle

3.03 3.19 3.09 3.26 2.33 2.76 2.37 2.43 2.33 2.79 2.39 2.90 2.56 3.03

Head width at level of

jaw angles

3.16 3.16 2.97 3.03 2.37 2.55 2.34 2.23 2.30 2.44 2.07 2.28 2.44 2.94

Head width at

anteriormost

nuchal collar

groove

3.20 3.36 3.08 3.08 2.31 2.59 2.36 2.36 2.28 2.44 2.41 2.10 2.63 3.01

Length of first nuchal

collar, measured

laterally

1.98 1.84 1.75 1.61 1.75 1.42 1.47 1.20 1.28 1.37 1.21 1.33 1.62 1.80

Length of second

nuchal collar,

measured laterally

2.21 2.06 2.02 2.14 1.93 1.62 1.55 1.60 1.75 1.52 1.38 1.37 1.54 1.88

Distance between

anterior margin of

upper lip and tip of

snout

1.22 1.10 1.12 1.10 0.80 1.00 0.96 0.87 0.91 0.96 0.96 0.88 1.12 1.00


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Supplementary (Continued)


Distance between

tentacle and naris

1.48 1.40 1.65 1.63 1.00 0.96 1.07 0.93 0.96 0.93 1.08 0.88 0.82 1.39

Distance between

tentacles

2.56 2.56 2.23 2.36 1.78 1.83 1.91 1.68 1.67 1.91 1.61 1.59 1.90 2.26

Distance between

nares

1.37 1.37 1.21 1.15 1.02 0.85 1.08 1.02 0.95 1.25 0.87 1.07 1.15 1.18

Distance betweennaris and jaw angle

3.63 3.74 3.55 4.02 2.88 3.27 2.69 2.82 3.15 2.66 2.47 3.19 2.71 3.09

Distance between

naris and tip of

snout

0.65 0.67 0.59 0.51 0.47 0.49 0.55 0.43 0.42 0.45 0.47 0.40 0.54 0.59

Distance between

naris and margin of

upper lip

0.95 1.28 1.14 0.93 1.01 0.74 0.73 0.72 0.90 0.98 0.81 0.80 0.98 0.88

Distance between

tentacle and jaw

angle

1.65 1.88 1.80 2.18 1.41 1.67 1.50 1.47 1.60 1.40 1.46 1.59 1.57 1.91

Distance between

tentacle and tip of

snout

1.79 2.03 2.09 2.12 1.45 1.55 1.63 1.41 1.52 1.56 1.38 1.43 1.66 1.81

Distance between

tentacle and margin

of upper lip

0.40 0.36 0.46 0.30 0.48 0.24 0.24 0.24 0.22 0.20 0.23 0.18 0.33 0.45

Width at midbody 3.57 4.50 3.50 3.92 3.01 2.61 2.88 2.50 2.71 2.93 2.36 2.39 3.28 3.42

Width at anterior

margin of terminal

shield

3.51 3.74 3.50 4.22 2.52 2.58 2.50 2.24 2.51 2.42 2.21 2.57 3.32 3.69


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Supplementary (Continued)


Length of terminal

shield, measured

laterally

3.41 3.44 2.81 3.48 3.34 3.16 2.23 2.30 2.14 2.01 2.29 2.19 2.54 3.24

Circumference at

midbody

13.10 12.40 10.20 11.60 9.30 9.80 8.00 7.70 8.50 8.90 7.50 8.60 10.80 11.50

Circumference 5

annuli anterior to

terminal shield

11.50 10.40 9.60 10.80 8.80 8.30 7.80 7.40 8.20 7.60 7.30 7.90 9.40 10.70

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